Apples

1.1.
1.4.
1.2.
1.5.
1.3.
Plate 1.1. Malus sieversii is a predominant overstorey species in the forests of eastern Kazakhstan.
Plate 1.2. Morphological variation among fruit collected in eastern Kazakhstan from trees of Malus sieversii.
Plate 1.3. Several species of Malus are frequently used as ornamental trees. Here different crab apple
selections make an attractive avenue.
Plate 1.4. A selection of over 50 older apple cultivars. Included in those shown are ‘Glory of England’, ‘Grimes
Golden’, ‘Lady Hamilton’, ‘Nonpareil’, ‘Rhode Island Greening’ and ‘Winter Banana’.
Plate 1.5. Very high-density (over 4000 trees ha-1) super-spindle apple trees growing near Bolzano in the Alto
Adige region of northern Italy. Nursery stock is being grown between the rows.
1.6. 1.7.
1.8.
1.10.
1.9.
Plate 1.6. High-density slender-spindle apple trees, typically planted with 3.0–3.5 m between rows and 1.0–1.25 m
within the rows, growing in The Netherlands.
Plate 1.7. Semi-intensive ‘Royal Gala’ apples, trained as centre-leader trees and typically grown with 5 m x 3 m
spacings, in Hawke’s Bay, New Zealand. Moderate year-round conditions, high sunshine hours and deep fertile
soils make this one of the most productive apple-producing regions of the world.
Plate 1.8. Semi-intensive centre-leader-trained apple trees growing near Grabouw in South Africa. Hot summer
conditions and mild winters allow the commercial production of both apples (foreground) and citrus
(background) in the same area.
Plate 1.9. Large ‘Rome Beauty’ trees on seedling rootstock, planted with 7.5 m x 7.5 m spacings in Ohio,
exemplify older plantings in the USA.
Plate 1.10. Extensive apple plantings, primarily of ‘Red Delicious’, trained as centre-leader trees, in
Washington State, USA. High summer and cold winter temperatures typify the growing conditions in this arid,
continental region.
3.1.
1.11.
3.2.
3.4.
3.3.
Plate 1.11. Production in China, now the world’s largest producer of apples (20 million t in 2001). These
5-year-old ‘Fuji’ trees on Malus prunifolia rootstock, are planted at 4 m x 3 m spacing. Vegetables are being
cultivated between the rows.
Plate 3.1. Modern breeding programmes generate many hundreds of progeny from each cross between selected
parents. Large land areas are needed to accommodate the seedling populations, which, because of the need to
grow through a juvenile phase, may need to be maintained for several years before the onset of flowering and the
initial evaluation of potential value.
Plate 3.2. Progeny arising from one cross can produce a very diverse range of fruit types – including different
sizes, colours and shapes.
Plate 3.3. ‘Pacific Rose’.
Plate 3.4. ‘Sci’ fresh apples now being marketed as ‘Jazz’.
3.5. 4.1.
4.3.
4.2.
4.4.
Plate 3.5. Colour sports of ‘Gala’ developed naturally as mutations.

Plate 4.1. ‘Delicious’, unspecified red strain (from Bruce Barritt).
Plate 4.2. ‘Golden Delicious’, Reinders strain (from Praktijkonderzoek Plant and Omgeving, The Netherlands).
Plate 4.3. ‘Fuji’ (from Agriculture and Agri-Food Canada).
Plate 4.4. ‘Granny Smith’ (from Bruce Barritt).
4.5. 4.6.
4.7.
4.9.
4.8.
Plate 4.5. ‘Imperial Gala’ (from Bruce Barritt).

Plate 4.6. ‘Jonathan’ (from Praktijkonderzoek Plant and Omgeving, The Netherlands).
Plate 4.7. ‘Jonagold’ (from Praktijkonderzoek Plant and Omgeving, The Netherlands).
Plate 4.8. ‘McIntosh’ (from Bruce Barritt).
Plate 4.9. ‘Rome Beauty’, unspecified red strain (from Agriculture and Agri-Food Canada).
4.11
4.10.
5.2.
4.12.
5.1.
Plate 4.10. ‘Braeburn’ (from Praktijkonderzoek Plant and Omgeving, The Netherlands).
Plate 4.11. ‘Elstar’ (from Bruce Barritt).
Plate 4.12. ‘Cox’s Orange Pippin’ (from Bruce Barritt).
Plate 5.1. Bed system of apple scions growing on M.27 rootstock in the UK.
Plate 5.2. A typical three-row bed system of ‘Queen Cox’ apples planted on M.9 EMLA rootstocks in the UK.
5.3.
6.3.
5.4.
6.1. 6.2.
Plate 5.3. Two-row system of seventh-leaf ‘Granny Smith’ on M.26 rootstock in the USA.
Plate 5.4. Traditional vigorous 40-year-old trees of the cultivar ‘Early Victoria’ on seedling rootstocks.
Plate 6.1. Obliquely planted M.9 liners as starting material for a new layer bed.
Plate 6.2. Example of a machine for harvesting rooted plants from a layer bed.
Plate 6.3. Layer bed of M.9: at the right, first season: at the left, second season.
6.4. 6.5.
6.6.
6.7.
6.8.
Plate 6.4. Mature layer bed of M.9.

Plate 6.5. Bundles of rootstock hardwood cuttings harvested after bedding for 1 year in the nursery.
Plate 6.6. Hard-pruned hedges of apple rootstocks.
Plate 6.7. In vitro micropropagated apple shoots prior to subculturing.
Plate 6.8. ‘Starkspur Golden Delicious’, rooted in vitro.
6.9. 6.11.
6.10.
6.12.
6.13.
Plate 6.9. Ex-micropropagated ‘Starkspur Golden Delicious’ in the orchard 4 years after being planted as a small
whip. Note the lack of fruits.
Plate 6.10. One-year-old trees of ‘Red Boskoop’ on M.9. Trees from left to right: unsprayed, or sprayed eight times
(weekly) with 50, 100, 300 or 600 p.p.m. benzyladenine, respectively, to improve feathering.
Plate 6.11. ‘Snip’ tree of ‘Elstar’ on M.9. Tree made from a table graft and cut back at 50 cm after the first nursery
year.
Plate 6.12. Productive interstem tree of ‘Elstar’ on M.9 in the second leaf in the orchard.
Plate 6.13. Chip budding.
9.1.
6.14.
9.2.
8.1.
9.3.
Plate 6.14. Tying the chip to the rootstock.

Plate 8.1. Detailed monitoring of soil-moisture status and tree transpiration rates in experimental systems improves
knowledge of apple tree water use. Enhanced water-use efficiency through better irrigation scheduling can result
from such studies.
Plate 9.1. Whole-tree gas-exchange chambers used for determined photosynthesis rates of intact apple canopies.
Impacts of factors such as water stress and crop load can be assessed through using such sophisticated
equipment.
Plate 9.2. Fish-eye photograph used to assess canopy openness and light penetration as influenced by training
system or pruning.
Plate 9.3. Reflective mulch used under ‘Fuji’ trees to improve fruit colour.
9.4. 10.1.
9.5.
11.2.
11.1.
Plate 9.4. ‘Fuji’ trees with paper bags around individual fruit to enhance colour when bags are removed (from
Bruce Barritt).
Plate 9.5. ‘Fuji’ apples previously bagged with stems clipped and ready for market. Bagging normally enhances
value as fruit are sold individually (from Bruce Barritt).
Plate 10.1. Large controlled environment rooms being used to study the impacts of temperature on apple fruit
growth rates at different times throughout the growing season.
Plate 11.1. Orchard in Washington State, USA, planted close to a lake to reduce risk of spring frost and as a
water-supply.
Plate 11.2. Top. ‘Frost ring’ on ‘Gala’ apple resulting from cold injury to the surface cells within a few days
following full bloom. The injured cells callused, giving the surface a russeted appearance. Bottom. ‘Delicious’ and
‘Golden Delicious’ apples in which the outer flesh were frozen approximately 3 weeks after full bloom. The skin
split but the cracks callused and healed. In neither case were the temperatures sufficiently cold for a long enough
period to freeze the developing seeds; therefore the fruit matured, although severely damaged by the cold.
11.3. 11.4.
11.5.
12.1.
12.2.
Plate 11.3. Orchard in British Columbia, Canada, planted across a hillside.

Plate 11.4. Orchard planted on the contour of the hillside.
Plate 11.5. Young trees in British Columbia, Canada, in a modern orchard trained to the axis system.
Plate 12.1. Leaf symptoms of potassium deficiency on ‘McIntosh’.
Plate 12.2. Leaf symptoms of magnesium definiency.
12.4.
12.3.
12.5.
12.7.
12.6.
Plate 12.3. Bitter pit – a calcium deficiency disorder – as manifested in ‘Golden Delicious’ apple.
Plate 12.4. Internal breakdown of ‘Spartan’ caused by calcium deficiency.
Plate 12.5. Boron deficiency in apple as manifested by drying and shrivelling of blossom.
Plate 12.6. Boron deficiency causing surface cracking of fruit.
Plate 12.7. Boron toxicity on ‘Golden Delicious’.
12.8. 12.9.
12.10.
13.1. 13.2.
Plate 12.8. Zinc-deficiency symptoms manifested on young ‘Delicious’/M.9 apple trees.

Plate 12.9. Iron deficiency.
Plate 12.10. Manganese deficiency.
Plate 13.1. Colour and size differences of fruit from trees growing in mowed sod grass (right) compared with
bare-soil residual-herbicide treatments (left), on the same harvest date for ‘Jonagold’ apples in a New York orchard.
Cumulative fruit yields (1989–1994) were about 25% lower in the sod-grass treatment, but fruit quality and market
value were consistently higher in sod than in the herbicide treatment (from Merwin and Stiles, 1994).
Plate 13.2. Soil-surface and ground-cover-vegetation conditions beneath trees during early December 2000, in
contiguous orchard-floor management (OFM) plots after 8 years of treatments with pre-emergence herbicides
(bare soil on left) versus a post-emergence herbicide (moss- and weed-covered soil on right) in a New York
orchard. Despite the substantial ‘weed’ ground cover during dormant seasons in the post-emergence herbicide
treatment, cumulative yields (1994–2000) were greater in this OFM system compared with bare-soil residual
herbicides. Note that weathering has eroded the upper 1 cm topsoil of fine soil particles, exposing a gravelly layer
in the residual-herbicide treatment (from an ongoing study described in Merwin et al., 1996).
13.5.
13.3.
14.2.
13.4.
14.1.
Plate 13.3. A front-mounted shielded-boom herbicide sprayer (Phil Brown Welding Co., Conklin, Michigan, USA)
for applications within the tree row. Metal or plastic shielding above nozzles reduces drift and spray damage to
tree trunks and foliage, facilitating herbicide application under moderately windy conditions.
Plate 13.4. Orchard mowers with low vertical profiles and adjustable side-wings permit close and effective mowing
of various alley widths and within tree rows without damage to low hanging branches laden with fruit around har-
vest time (mower shown is PerfectTM model DR365, as manufactured by Van Wamel, BV, BenedenLeeuwen, The
Netherlands).
Plate 13.5. Top and bottom: views of a prototype shrouded propane flame weeder developed by I. Merwin, J. Ray
and K. Bittner at Cornell University, for weed suppression beneath trees and vines. The unit floats on two ground
skids, with dual torches facing forward beneath a protective metal shroud, which reduces fuel consumption and
minimizes heat damage to trees, vines and trickle-irrigation lines.
Plate 14.1. Large apple trees on seedling rootstock with minimal pruning and training.
Plate 14.2. Unpruned (left) and summer-pruned container-grown apple trees showing the reduction in shoot
growth, leaf size and root growth (from Taylor and Ferree, 1981).
14.3. 14.4.
14.5.
14.6.
15.1.
Plate 14.3. Mechanical root pruning of apple trees at bloom.

Plate 14.4. Apple tree trained in an ornamental form using bending and pruning.
Plate 14.5. Chain-saw cut used to interrupt phloem transport and reduce apple tree growth (from Steve Hoying).
Plate 14.6. Plastic sleeves used to induce bud break on vigorous young apple trees (from Steve Hoying).
Plate 15.1. Traditional globe-shaped apple tree of ‘McIntosh’ on seedling rootstock.
15.2. 15.3.
15.4.
15.6.
15.5.
Plate 15.2. Central-leader apple tree of ‘Gala’ on MM.106 rootstock with four distinct tiers of branches.
Plate 15.3. Slender-spindle apple tree of ‘Gala’ on M.9 rootstock with leader zigzagging to limit tree height (from
Bruce Barritt).
Plate 15.4. Vertical-axis apple trees of ‘Gala’ on M.9 rootstock (from Bruce Barritt).
Plate 15.5. Vertical-axis apple orchard of ‘Golden Delicious’ on M.9 rootstock. Upper arrow denotes large branch to
be removed back to an angled stub. Lower arrow denotes renewal branch arising from the stub of a previous year's
pruning cut.
Plate 15.6. Solaxe apple tree of ‘Jonagold’ on M.9 rootstock with lower branches bent down in a pendant position.
15.7.
15.8.
15.9.
15.11.
15.10.
Plate 15.7. Slender-pyramid apple tree of ‘Gala’ on M.26 rootstock with distinct tiers of horizontal branches.
Plate 15.8. HYTEC apple trees of ‘Gala’ on M.9 rootstock with the central leader tied over at a 45 o angle to reduce
the vigour of the top.
Plate 15.9. Super-spindle apple orchard of ‘Empire’ on M.9 rootstock.
Plate 15.10. Horizontal-palmette-trellis tree of ‘Jonagold’ on Mark rootstock.
Plate 15.11. Lincoln-canopy apple orchard of ‘Delicious’ on MM.106 rootstock with vigorous shoot growth arising
from the horizontal canopy.
15.12. 15.13.
15.14.
15.15.
16.1.
Plate 15.12. Geneva Y-trellis apple orchard of ‘Empire’ on M.26 rootstock.

Plate 15.13. MIA trellis (A-shaped trellis) apple orchard of ‘Gala’ on MM.106 rootstock.
Plate 15.14. Gütingen V-slender-spindle apple orchard of ‘Jonagold’ on M.9 rootstock.
Plate 15.15. V-super-spindle apple orchard of ‘Gala’ on M.9 rootstock.
Plate 16.1. ‘Redchief Delicious’ apple injury caused by Carbaryl + Accel + Regulaid under certain environmental
conditions: (A) 2 weeks after treatment; (B) near harvest; and (C) at harvest (from Byers et al., 2000a).
18.1. 18.2.
18.4.
18.3.
18.5.
Plate 18.1. Dieback of apple spurs and shoots caused by fire blight, the disease caused by the bacterium
Erwinia amylovora (from A.L. Jones).
Plate 18.2. Apple spur and shoot with fire blight. Infected foliage appears scorched. The ‘shepherd’s crook’
recurving of the shoot tip is a diagnostic symptom of the disease (from A.L. Jones).
Plate 18.3. Apple fruit with fire blight, note ooze (from A.L. Jones).
Plate 18.4. ‘McIntosh’ apple leaf with scab, caused by Venturia inaequalis (from A.L. Jones).
Plate 18.5. Symptoms of apple scab on immature ‘McIntosh’ fruit (from A.L. Jones).
18.6. 18.7.
18.8.
18.9. 18.10.
Plate 18.6. Foliar/shoot symptoms of powdery mildew, caused by Podosphaera leucotricha (from A.L. Jones).
Plate 18.7. Net-russet symptoms of powdery mildew on infected apple fruit (from A.L. Jones).
Plate 18.8. European brown rot on apple fruit (from A.L. Jones).
Plate 18.9. Depressed bitter-rot lesions on apple, caused by Colletotrichum gloeosporioides or Colletotrichum
acutatum (from A.L. Jones).
Plate 18.10. Cortland apples with black rot; one is mummified with pycnidia. The disease is caused by
Botryosphaera obtusa (from A.L. Jones).
18.11. 18.12.
18.13.
18.14.
18.15.
Plate 18.11. Bot (white) rot of apple, caused by Botryosphaeria dothidea. Several ‘Golden Delicious’ with white rot.
Apple lower left with black rot (from A.L. Jones).
Plate 18.12. Sooty blotch and a few fly-speck lesions on apple (from A.L. Jones).
Plate 18.13. Black, shiny fly-speck lesions on apple, caused by Schizothyrium pomi. Sooty blotch is evident above
fly-speck lesions (from A.L. Jones).
Plate 18.14. Brooks spot at the calyx end of ‘Grimes Golden’ apple, caused by Mycosphaerella pomi (from A.L.
Jones).
Plate 18.15. Purple spots on apple leaf caused by Brooks spot (from A.L. Jones).
18.16.
18.17.
18.18.
18.20.
18.19.
Plate 18.16. Alternaria blotch on leaves of ‘Delicious’, caused by Alternaria mali (from A.L. Jones).
Plate 18.17. Small black spot of black pox on ‘Grimes Golden’ apple. The disease is caused by Helminthosporium
papulosum (from A.L. Jones).
Plate 18.18. Necrotic leaf blotch, a physiological disorder, on cultivar ‘Golden Delicious’ (from A.L. Jones).
Plate 18.19. Phytophthora crown and root rot on apple rootstock, caused by fungi in the genus Phytophthora
(from A.L. Jones).
Plate 18.20. European canker, caused by Nectria galligena. Zonate cankers are characteristic of the disease
(from A.L. Jones).
18.21. 18.22.
18.24.
18.23. 18.25.
Plate 18.21. Blue mould, caused by Penicillium expansum, on ‘Golden Delicious’ apple (from A.L. Jones).
Plate 18.22. Flat apple on cultivar ‘Red Delicious’. The virus causing the disease (cherry rasp-leaf virus) is
vectored by the nematode Xiphenema americanum (from W.E. Howell).
Plate 18.23. Apple mosaic on cultivar ‘Golden Delicious’ (from W.E. Howell).
Plate 18.24. Dapple apple on ‘Bisbee Red Delicious’, note concentration of spotting near the calyx (from K.C.
Eastwell).
Plate 18.25. Apple scar skin. This disease is caused by a viroid (from W.E. Howell).
18.26. 19.2.
19.3.
19.1.
19.4.
Plate 18.26. Apple green-crinkle symptoms on cultivar ‘Golden Delicious’ (from W.E. Howell).
Plate 19.1. Adult lygus (Lygus lineolaris Palisot de Beauvois) stings apple fruitlets, but does not reproduce on
apple.
Plate 19.2. Thrips damage (pansy spot) to ‘Delicious’ (left) and ‘Granny Smith’ apple. The damage on ‘Delicious’
will colour over by harvest, but the damage on ‘Granny Smith’ will still be visible.
Plate 19.3. Codling moth (Cydia pomonella) damage to apple; fully grown larva feeding in the core.
Plate 19.4. Adult oriental fruit moth (Grapholita molesta (Busck)); larvae are internal fruit feeders.
19.5. 19.6.
19.8.
19.7.
19.9.
Plate 19.5. Apple maggot (Rhagoletis pomonella) adult fly with characteristic wing-banding pattern.
Plate 19.6. Ectoparasitic larvae of the eulophid wasp Colpoclypeus florus Walker attacking the larva of oblique-
banded leaf-roller, Choristoneura rosaceana (Harris).
Plate 19.7. Adult oblique-banded leaf-roller, Choristoneura rosaceana (Harris).
Plate 19.8. European red-mite adult females. White spots are bases of large dorsal setae.
Plate 19.9. Two-spotted spider mite (Tetranychus urticae Koch) adult female. This extremely polyphagous species
has a cosmopolitan distribution.
19.10. 19.11.
19.13.
19.12.
20.1. 20.2.
Plate 19.10. Typhlodromus (Galandromus) occidentalis, the principal mite predator in the arid growing regions of
the western USA.
Plate 19.11. The San José scale, Quadraspidiotus perniciosus (Comstock), attacks both shoots and fruit.
Plate 19.12. Apple aphids, Aphis pomi De Geer, infest shoots and leaves, removing plant phloem.
Plate 19.13. Woolly apple aphid, Eriosoma lanigerum (Hausmann), causes galls on twigs (in the leaf axils) and
roots.
Plates 20.1 and 20.2. Pruning during early to mid-endodormancy can be fatal. The trees in Plate 20.1 were not
pruned. The trees in Plate 20.2 were pruned in early November (northern hemisphere) and were dead by the
following July. The photos were taken from the same location in late May and show peach trees but apples can
also be affected.
20.5.
20.3.
20.6.
20.4.
20.7.
Plate 20.3. Towers used to monitor inversions and wind-machine effects.

Plate 20.4. Heater plumes.
Plate 20.5. Under-tree sprinkling for freeze protection.
Plate 20.6. Wind machines in a high mountain valley.
Plate 20.7. Over-tree sprinkling for apple-bloom delay. Delayed trees (still nearly dormant) in the background are
white from calcium carbonate deposition. Controls, in the foreground, are nearing first bloom.
21.1. 21.2.
22.1A.
22.1B. 22.1C.
Plate 21.1. A large air-blast sprayer in operation. New methods use improved equipment with more targeted
application, less persistent chemicals and timing based on economic thresholds rather than calendar-based intervals.
Plate 21.2. Pheromone traps are used for a number of purposes including detection of adult male insects,
determining the efficacy of mating disruption and for arriving at spray-application thresholds.
Plate 22.1. Orchards of dwarfing trees can incorporate many practical options for increasing biological diversity
and thus enhancing biological stability. (A) Strips sown in wild flowers and herbs; (B) nesting box for tits and
earwig nest; (C) nesting block for wild bees.
22.2A.
22.2B.
22.2C.
22.2D.
22.2E.
Plate 22.2. Soil management – methods and tools. (A) The newly developed Ladurner mechanical hoe impresses
with a construction guaranteeing operational comfort and good performance even in difficult conditions. (B)
Undercutters (Müller RPM). Good on light soils. Problems occur in dense swards. (C) Crumbler with vertical
cutters (Humus-Planet). (D) Disc plough (Spedo). (E) Thermal weed control: this device combines heat treatment
with an open flame for weeding around the trunk and an infrared emitter for the strip.
22.2F. 22.2G.
22.2H.
22.3.
Plate 22.2. Continued (F) FiBL's (Forschungsinstitut für Biologischer Landbau) ‘sandwich system’ is still in the
development phase (ground-cover management, mechanical hoe for strip cultivation). It is designed to allow for the
use of inexpensive and efficient mechanical hoeing equipment. The low-growing, diverse herbal ley in the
middle of the in-row strip can be advantageous in supporting beneficials and in helping to maintain soil fertility.
(G) Mechanized laying of mulch sheeting. (H) Bark mulch applied with a mulch spreader.
Plate 22.3. As synthetic thinning sprays are not permitted, the development of the rope machine, a mechanical
tool for blossom thinning, has been a major step forward in solving one of the key problems of organic apple
production. This development also significantly improves the conditions for converting larger orchards to organic
production.
23.1.
23.2.
23.3.
24.1.
Plate 23.1. Fruit inspection in a modern packing-house.

Plate 23.2. Loading a ship for export from New Zealand in the 1940s. Wooden cases are being handled
individually at each stage of the transport process.
Plate 23.3. Loading a ship for export from New Zealand in the 1990s. Corrugated-cardboard cartons, which have
replaced wooden boxes, are stacked on fork-lift pallets for enhanced handling efficiency.
Plate 24.1. Array of processed apple products of Knouse Foods Cooperative Inc. (from Knouse Foods Cooperative
Inc.).
Apples - Chap 00 11/4/03 11:54 am Page i
Apples
Botany, Production and Uses

Apples - Chap 00 11/4/03 11:54 am Page ii
The editors dedicate this book to all the professors,

teachers, extension personnel, commercial growers
and other professionals who willingly shared
their knowledge, inspired their students and
contributed to the extensive knowledge that is
represented in this book.
Apples - Chap 00 11/4/03 11:54 am Page iii
Apples
Botany, Production and Uses
Edited by
D.C. Ferree
Department of Horticulture and Crop Science, Ohio State University, USA
and
I.J. Warrington
Department of Horticultural Science, Massey University, Palmerston North,

New Zealand
CABI Publishing
Apples - Chap 00 11/4/03 11:54 am Page iv
CABI Publishing is a division of CAB International

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Apples : botany, production, and uses / edited by D.C. Ferree and I.J.
Warrington
p. cm.
Includes bibliographical references.
ISBN 0-85199-592-6
1. Apples. 2. Apples--United States. I. Ferree, David C. (David
Curtis), 1943- II. Warrington, I. J. (Ian J.) III. Title.
SB363 .A67 2003
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2002013984
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Typeset in 9 on 11 pt Palatino by Columns Design Ltd, Reading

Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn
Apples - Chap 00 11/4/03 11:54 am Page v
Contents
Contributors vii
Preface ix
Acknowledgements xi
PART I: INTRODUCTION
1. Taxonomic Classification and Brief History 1
J.J. Luby
2. World Production, Trade, Consumption and Economic Outlook for Apples 15
D. O’Rourke
PART II: PLANT MATERIALS

3. Genetic Improvement of Apple: Breeding, Markers, Mapping and Biotechnology 31
S.K. Brown and K.E. Maloney
4. Characteristics of Important Commercial Apple Cultivars 61
C.R. Hampson and H. Kemp
5. Apple Rootstocks 91
A.D. Webster and S.J. Wertheim
6. Propagation and Nursery Tree Quality 125
S.J. Wertheim and A.D. Webster
PART III: APPLE PHYSIOLOGY AND ENVIRONMENTAL INFLUENCES

7. Flowering, Pollination and Fruit Set and Development 153
F. Dennis, Jr
8. Water Relations of Apples 167
A.N. Lakso
9. Light Relations 195
L. Corelli Grappadelli
10. Temperature 217
J.W. Palmer, J.P. Privé and D.S. Tustin
PART IV: ORCHARD AND TREE MANAGEMENT

11. Selecting the Orchard Site, Site Preparation and Orchard Planning and
Establishment 237
J.A. Barden and G.H. Neilsen
12. Nutritional Requirements of Apple 267
G.H. Neilsen and D. Neilsen
v
Apples - Chap 00 11/4/03 11:54 am Page vi
vi Contents
13. Orchard-floor Management Systems 303

I.A. Merwin
14. Pruning and Training Physiology 319
D.C. Ferree and J.R. Schupp
15. Apple-orchard Planting Systems 345
T.L. Robinson
16. Flower and Fruit Thinning and Vegetative : Fruiting Balance 409
R.E. Byers
17. Endogenous Hormones and Bioregulator Use on Apples 437
D.W. Greene
PART V: CROP PROTECTION

18. Diseases of Apple 459
G.G. Grove, K.C. Eastwell, A.L. Jones and T.B. Sutton
19. Ecology and Management of Apple Arthropod Pests 489
E.H. Beers, D.M. Suckling, R.J. Prokopy and J. Avilla
20. Apple-orchard Freeze Protection 521
S.D. Seeley and J.L. Anderson
21. Integrated Fruit Production for Apples – Principles and Guidelines 539
J. Avilla and H. Riedl
22. Organic Apple Production – with Emphasis on European Experiences 551
F. Weibel and A. Häseli
PART VI: HARVESTING, HANDLING AND UTILIZATION

23. Principles and Practices of Postharvest Handling and Stress 585
C.B. Watkins
24. Production and Handling Techniques for Processing Apples 615
R.M. Crassweller and G.M. Greene, II
Index 635
Colour plate section after p. 436

Apples - Chap 00 11/4/03 11:54 am Page vii
Contributors
J. LaMar Anderson, Plants, Soils, and Biometeorology Department, Utah State University, Logan,
UT 84322, USA.
Jesús Avilla, Centro UdL-IRTA de R+D de Lleida, University of Lleida, Rovira Roure 177, 25198
Lleida, Spain.
John A. Barden, Department of Horticulture, Virginia Polytechnic Institute and State University,
Blacksburg, VA 24061, USA.
Elizabeth H. Beers, Washington State University, Tree Fruit Research and Extension Center, 1100
N. Western Avenue, Wenatchee, WA 98801, USA.
Susan K. Brown, Department of Horticultural Sciences, Cornell University, New York State
Agricultural Experiment Station, Geneva, NY 14456, USA.
Ross E. Byers, Department of Horticulture, Virginia Polytechnic Institute and State University,
Winchester, VA 22602, USA.
Luca Corelli Grappadelli, Dipartimento di Colture Arboree, University of Bologna, Via Fillippo Re.
6, 40126 Bologna, Italy.
Robert M. Crassweller, The Pennsylvania State University, Department of Horticulture, Fruit
Research and Extension Center, PO Box 330, Biglerville, PA 17307, USA.
Frank Dennis, Jr, Department of Horticulture, Michigan State University, East Lansing, MI 48824,
USA.
Kenneth C. Eastwell, Irrigated Agriculture Research and Extension Center, Washington State
University, Prosser, WA 99350, USA.
David C. Ferree, Department of Horticulture and Crop Science, Ohio Agricultural Research and
Development Center, Ohio State University, Wooster, OH 44691, USA.
Duane W. Greene, Department of Plant and Soil Sciences, University of Massachusetts, Box 30910,
Amherst, MA 01003, USA.
George M. Greene, II, The Pennsylvania State University, Department of Horticulture, Fruit
Research and Extension Center, PO Box 330, Biglerville, PA 17307, USA.
Gary G. Grove, Irrigated Agriculture Research and Extension Center, Washington State University,
Prosser, WA 99350, USA.
Cheryl R. Hampson, Pacific Agri-Food Research Center, Agriculture and Agri-Food Canada,
Summerland, BC V0H 1Z0, Canada.
Andreas Häseli, Research Institute of Organic Agriculture (Forschungsinstitut für biologischen
Landbau (FiBL)), Ackerstrasse, Postfach, CH-5070 Frick, Switzerland.
Alan L. Jones, Department of Botany and Plant Pathology, Michigan State University, East
Lansing, MI 48824, USA.
Henk Kemp, Applied Plant Research, Fruit Section, Lingewal 1, 6668 LA Randwijk, The
Netherlands.
vii
Apples - Chap 00 11/4/03 11:54 am Page viii
viii Contributors
Alan N. Lakso, Fruit Crop Physiology Program, Cornell University, Department of Horticultural
Sciences, New York State Agricultural Experiment Station, Geneva, NY 14456, USA.
James J. Luby, Department of Horticultural Sciences, University of Minnesota, 342 Alderman Hall,
1970 Folwell Avenue, St Paul, MN 55108, USA.
Kevin E. Maloney, Department of Horticultural Sciences, Cornell University, New York State
Agricultural Experiment Station, Geneva, NY 14456, USA.
Ian A. Merwin, Department of Horticulture, Cornell University, Ithaca, NY 14853, USA.
Denise Neilsen, Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre,
Summerland, British Columbia, V0H 1Z0, Canada.
Gerry H. Neilsen, Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre,
Summerland, British Columbia, V0H 1Z0, Canada.
Desmond O’Rourke, Belrose Inc., 1045 NE Creston Lane, Pullman, WA 99163, USA.
John W. Palmer, The Horticulture and Food Research Institute of New Zealand Ltd, Nelson
Research Centre, PO Box 220, Motueka, New Zealand.
Jean P. Privé, Agriculture and Agri-Food Canada, PO Box 667, Boutouche, New Brunswick,
Canada.
Ronald J. Prokopy, Department of Entomology, University of Massachusetts, Fernald Hall, Box
30910, Amherst, MA 01003, USA.
Helmut Riedl, Mid-Columbia Agricultural Research and Extension Center, Oregon State
University, 3005 Experiment Station Drive, Hood River, OR 97031, USA.
Terence L. Robinson, Department of Horticultural Sciences, New York State Agricultural
Experiment Station, Cornell University, Geneva, NY 14456, USA.
James R. Schupp, Department of Horticultural Sciences, Cornell University, New York State
Agricultural Experiment Station, Hudson Valley Laboratory, Highland, NY 12528, USA.
Schuyler D. Seeley, Plants, Soils, and Biometeorology Department, Utah State University, Logan,
UT 84322, USA.
D. Max Suckling, The Horticulture and Food Research Institute of New Zealand Ltd, Gerald Street,
PO Box 51, Lincoln, Canterbury, New Zealand.
Turner B. Sutton, Department of Plant Pathology, North Carolina State University, Raleigh, NC
27695, USA.
D. Stuart Tustin, The Horticulture and Food Research Institute of New Zealand Ltd, Hawke’s Bay
Research Centre, Private Bag 1401, Havelock North, New Zealand.
Ian J. Warrington, Department of Horticultural Science, Massey University, Private Bag 11222,
Palmerston North, New Zealand.
Christopher B. Watkins, Department of Horticulture, Cornell University, Ithaca, NY 14853, USA.
Anthony D. Webster, Crop Science Department, Horticulture Research International, East Malling,
West Malling, Kent ME19 6BJ, UK.
Franco Weibel, Research Institute of Organic Agriculture (Forschungsinstitut für biologischen
Landbau (FiBL)), Ackerstrasse, Postfach, CH-5070 Frick, Switzerland.
S.J. Wertheim, Fruit Research Station, Lingewal 1, 6668 Randwijk, The Netherlands.
Apples - Chap 00 11/4/03 11:54 am Page ix
Preface
There is no fruit in temperate climates so universally esteemed, and so extensively cultivated, nor is
there any which is so closely identified with the social habits of the human species as the apple.
(Dr Robert Hogg, The Apple, 1851)
Although the precise origin of today’s apple is not entirely clear, it probably evolved from exten-
sive forests of apples in central Asia, particularly in Kazakhstan. Due to its unique qualities, peo-
ple collected and spread the most desirable types. Remains of apple have been reported in
historic sites dated to 6500 BC. Long-distance trade routes between the Mediterranean area and
various areas of Asia developed as early as 3500 BC and fostered the spread of both fresh and
dried apples. Theophrastus (around 320 BC) studied apples brought back to Greece from con-
quests of Alexander the Great. He described grafting and general tree care and also dwarf types
that later were used as rootstocks. Followers of both the Christian and Islam religions were
instrumental in the spread of apples throughout Europe, Africa and the New World. By 1826, the
Royal Horticultural Society of England had identified 1200 apple cultivars.
Commercial production of apples started as complements in gardens, as field borders or
as overstorey trees in pastures. Apples are produced commercially in most countries in the
temperate region of the world and also in some tropical areas with high altitude. In the last
100 years production has become increasingly intensified, with the use of dwarfing root-
stocks and training systems designed to improve orchard efficiency. Apple is unique among
fruit plants in having a range of rootstocks that permit development of a ‘designer tree size’
appropriate to the training system and management skills of modern orchardists.
In the last 50 years the development of herbicides, insecticides and fungicides has permit-
ted the production of high-quality fruit in many areas where production was previously diffi-
cult. Currently, as more information is gained through research, the trend is to reduce
pesticide inputs through integrated production systems or organic production. Apple breed-
ers are assisting by developing high-quality cultivars with resistances to the most serious
pests, through both conventional breeding and genetic engineering. Research in storage and
postharvest handling techniques have dramatically improved fruit quality and currently
apples are a quality product available throughout the year. Many of these current cultural
practices are based on research results of detailed studies of the effects of various aspects of
the environment on apple growth and development.
This book is an effort by 39 research scientists from eight countries to summarize the cur-
rent research information on apples in a comprehensive treatise. Authors attempted to provide
the information and physiology behind current cultural practices as well as future trends. The
objective was to provide horticultural students, research and extension personnel, professional
fruit growers and others with a comprehensive textbook on apples and their culture.
David C. Ferree
Ian J. Warrington
ix
Apples - Chap 00 11/4/03 11:54 am Page x
Apples - Chap 00 11/4/03 11:54 am Page xi
Acknowledgements
The editors and authors want to thank the following organizations for sponsoring the colour
section of this book:
The Horticulture and Food Research Institute of New Zealand Ltd; The New Zealand
Fruitgrowers Charitable Trust; The Ohio Fruit Growers Society; and The Ohio Fruit
Growers Marketing Association.
xi
Apples - Chap 00 11/4/03 11:54 am Page xii
Apples - Chap 01 11/4/03 11:00 am Page 1
1 Taxonomic Classification and Brief History
James J. Luby
Department of Horticultural Sciences, University of Minnesota, St Paul,
Minnesota, USA
1.1 The Origin and Spread of the Domesticated Apple 1

1.2 Taxonomy and Evolution 9
1.2.1 Apples in the family Rosaceae 9
1.2.2 Species in the genus Malus 11
1.1 The Origin and Spread of the believed it originated as a hybrid derived
Domesticated Apple from M. sylvestris Mill., Malus dasyphyllus
Borkh. (a synonym for M. pumila) and Malus
The common domesticated apple is puta- praecox Borkh. (a synonym for M. sylvestris
tively an interspecific hybrid complex, usu- var. praecox (Pall.) Ponomar.) (Korban and
ally designated Malus × domestica Borkh. Skirvin, 1984). Currently, however, Malus
(Korban and Skirvin, 1984) or M. domestica sieversii (Ledeb.) Roem. is hypothesized as
Borkh. (Phipps et al., 1990). Other synonyms, the key species in its origin (Ponomarenko,
now considered illegitimate, have been 1983; Vavilov, 1987; Roach, 1988; Way et al.,
applied, including Pyrus malus L., Malus malus 1990; Hokanson et al., 1997; Juniper et al.,
Britt., Malus pumila Mill. and Malus sylvestris 1998). M. sieversii is widespread in the
Mill. M. × domestica is now cultivated widely mountains of central Asia at elevations
in temperate latitudes or at high elevations in between approximately 1200 and 1800 m.
the tropics on all continents except Antarctica. The forests are extensive and M. sieversii is
The fruits are eaten fresh, dried or tinned or the dominant overstorey species in many
processed into juice, preserves or alcoholic areas (Plate 1.1). The fruit of M. sieversii is
beverages. Besides M. × domestica, fruits of highly variable (Plate 1.2) and individual
several other species are consumed fresh or trees resembling M. × domestica are com-
processed or are used for medicinal purposes monly found in the forests of this region but
and the plants are used as rootstocks (Table their precise history is difficult to ascertain.
1.1). Many species and interspecific hybrids Humans have inhabited and practised
are used as ornamental plants. nomadic agriculture in this region for thou-
The origin and ancestry of the M. × domes- sands of years. People of this region today
tica complex remain unknown. Borkhausen, will save desirable trees when the forest is
when first describing M. × domestica in 1803, cleared for agriculture (Ponomarenko, 1983)
© CAB International 2003. Apples: Botany, Production and Uses

(eds D.C. Ferree and I.J. Warrington) 1
Apples - Chap 01
Table 1.1. Malus species, synonyms and infraspecific classifications, from the taxonomy database of the US Department of Agriculture Germplasm Resources
2
Information Network (USDA, ARS, National Genetic Resources Program. Germplasm Resources Information Network (GRIN), 2000) and their chromosome
number, presence of apomixis, distribution and uses (from GRIN and also Phipps et al., 1990; Way et al., 1990; Zhang et al., 1993; Deng et al., 1995; Schuster
and Büttner 1995; Zhou, 1999).
11/4/03
Chromosome
number and Synonyms and infraspecific classifications and
Species apomixis (A) [putative origin of secondary species] Distribution Uses
11:00 am
Primary species
M. angustifolia (Aiton) Michx. 34 Eastern USA Ornamental, preserves
M. baccata (L.) Borkh. 34, A M. baccata var. baccata North-eastern China, eastern Ornamental, rootstock
M. rockii Rehder Siberia, Mongolia, northern
M. sibirica (Maxim.) Kom., nom. illeg. India, Bhutan, Nepal
Page 2
M. baccata f. gracilis Rehder
M. baccata f. jackii Rehder
M. baccata subsp. himalaica (Maxim.) Likhonos
J.J. Luby
M. baccata var. himalaica (Maxim.) C.K. Schneid.
M. baccata var. sibirica C.K. Schneid.
M. baoshanensis G.T. Deng – South-central China Rootstock
M. brevipes (Rehder) Rehder 34 Only known in cultivation Ornamental
M. coronaria (L.) Mill. 51, 68, A M. coronaria var. dasycalyx Rehder Eastern USA and Canada Ornamental, fruit, preserves
M. fragrans Rehder
M. glabrata Rehder
M. glaucescens Rehder
M. lancifolia Rehder
M. bracteata Rehder
M. daochengensis C.L. Li – South-central China
M. × domestica Borkh. 34, 51, 68, A M. malus (L.) Britton, nom. inval. Cultivated and naturalized in Fruit, preserves, beverage
M. pumila auct. temperate regions base, medicinal
M. sylvestris auct.
M. sylvestris var. domestica (Borkh.) Mansf.
M. doumeri (Bois) A. Chev – M. formosana Kawak. & Koidz. South-east China, Taiwan, Preserves
M. laosensis (Cardot) A. Chev. South-east Asia
M. florentina (Zuccagni) 34 M. crataegifolia (Savi) Koehne Turkey, Greece, Italy and Ornamental
C.K. Schneid. Balkans
Apples - Chap 01
M. floribunda Siebold ex 34 Only known in cultivation Ornamental
Van Houtte
M. fusca (Raf.) C.K. Schneid. 34 M. fusca var. diversifolia (Bong.) C.K. Schneid. Western USA and Canada
M. rivularis Douglas ex Hook.
11/4/03
M. diversifolia (Bong.) M. Roem.
M. halliana Koehne 34, 51 Central and eastern China, Ornamental, rootstock
Japan
M. honanensis Rehder – North-central China
11:00 am
M. hupehensis (Pamp.) 34, 51, 68, A M. theifera Rehder Central and south-east China Ornamental, rootstock, fruit,
Rehder beverage base, medicinal
M. ioensis (A.W. Wood) Britton 34, 51 M. ioensis var. texana Rehder Central USA Ornamental, fruit
M. jinxianensis J.Q. Deng & – Northern China
J.Y. Hong
Taxonomic Classification and History
Page 3
M. kansuensis (Batalin) 34 Central China
C.K. Schneid.
M. komarovii (Sarg.) Rehder – North-east China
M. leiocalyca S.Z. Huang – South-east China
M. maerkangensis – Central China
M.H. Cheng et al.
M. mandshurica (Maxim.) 34 M. sachalinensis Juz. Central and north-east China, Rootstock
Kom. M. baccata var. cerasifera (Spach) Koidz. far-eastern Russian, Japan
M. baccata var. mandshurica (Maxim.)
C.K. Schneid.
M. cerasifera Spach
M. mandshurica var.sachalinensis (Juz.) Ponomar.
M. melliana (Hand.-Mazz.) – South-east China Fruit, beverage base
Rehder
M. micromalus Makino 34, 51 Central and eastern China, Rootstock, fruit, medicinal
Japan
M. muliensis T.C. Ku – Central China
M. ombrophila Hand.-Mazz – South-central China
M. orientalis Uglitzk. – M. sylvestris subsp. orientalis (Uglitzk.) Browicz Caucasus, Iran
M. orthocarpa Lavallee ex – An uncertain taxon Only known in cultivation Ornamental
anon.
M. prattii (Hemsl.) 34 M. kaido Dippel Central China Fruit
C.K. Schneid.
3
Continued
Apples - Chap 01
4
Table 1.1. Continued.
Chromosome
11/4/03
M. prunifolia (Willd.) Borkh. 34 Central and eastern China Ornamental, rootstock

M. pumila Mill. 34 M. pumila var.niedzwetzkyana (Dieck) Eastern Europe Ornamental, rootstock
C.K. Schneid.
11:00 am
M. sylvestris var. niedzwetzkyana (Dieck)
L.H. Bailey
M. niedzwetzkyana Dieck
M. paradisiaca (L.) Medik.
M. dasyphylla Borkh.
Page 4
M. pumila var.paradisiaca (L.) C.K. Schneid.
M. sargentii Rehder, 34, 51, 68, A Only known in cultivation Ornamental
M. sieversii (Ledeb.) M. Roem. – M. sieversii subsp. turkmenorum (Juz. & Popov) Central Asia Rootstock, fruit, preserves
Likhonos
J.J. Luby
M. sieversii var. turkmenorum (Juz. & Popov)
Ponomar.
M. sieversii var. kirghisorum (Al. Fed. & Fed.)
Ponomar.
M. kirghisorum Al. Fed. & Fed.
M. turkmenorum Juz. & Popov
M. sikkimensis (Wenz.) 51, A South-central China, northern Rootstock, ornamental
Koehne ex C.K. Schneid. India, Bhutan
M. spectabilis (Aiton) Borkh. 34, 51 Eastern China Ornamental
M. sylvestris Mill. – M. praecox (Pall.) Borkh. Europe Ornamental, fruit, preserves
M. sylvestris var. praecox (Pall.) Ponomar.
M. toringo (Siebold) Siebold 34, 51, A M. sieboldii (Regel) Rehder Eastern China, Japan, Korea Ornamental, rootstock
ex de Vriese M. sieboldii var. arborescens Rehder
M. toringoides (Rehder) 51, 68, A M. transitoria var. toringoides Rehder Central China Rootstock
Hughes
M. transitoria (Batalin) 34, 51 North central China Rootstock
C.K. Schneid.
M. tschonoskii (Maxim.) 34 Japan
C.K. Schneid.
Apples - Chap 01
M. xiaojinensis M.H. Cheng & – Central China Rootstock
N.G. Jiang
M. yunnanensis (Franch.) 34 M. yunnanensis var. veitchii (Veitch) Rehder South-central China Ornamental, rootstock
C.K. Schneid. M. yunnanensis var. yunnanensis
11/4/03
M. zumi (Matsum.) Rehder 34 M. zumi var. calocarpa (Rehder) Rehder Japan
M. sieboldii var. calocarpa Rehder
Docynia indica (Wall.) Decne. – M. docynioides C.K. Schneid. Eastern Himalayas,
south-east Asia
11:00 am
Eriolobus trilobata (Poir.) 34 M. trilobata (Poir.) C.K. Schneid. Eastern Mediterranean
M. Roem.
Secondary species
M. × adstringens Zabel 34, 51 [= M. baccata × M. pumila] Only cultivated Ornamental
M. × arnoldiana (Rehder) 34 [= M. baccata × M. floribunda] Only cultivated Ornamental
Taxonomic Classification and History

Sarg. ex Rehder M. floribunda var. arnoldiana Rehder
Page 5
M. × asiatica Nakai – [= M. prunifolia × M. sieversii] Cultivated in east Asia Rootstock, fruit, preserves
M. prunifolia var. rinkii (Koidz.) Rehder
M. ringo Siebold ex Carriere
M. × astracanica hort. ex – [= M. prunifolia × M. pumila] Only cultivated Ornamental
Dum. Cours.
M. × atrosanguinea (Spath) – [= M. halliana × M. toringo] Only cultivated Ornamental
C.K. Schneid.
M. × dawsoniana Rehder 34 [= M. domestica × M. fusca] Only cultivated Ornamental
M. × hartwigii Koehne 34 [= M. baccata × M. halliana] Only cultivated Ornamental
M. × magdeburgensis Hartwig – [= M. pumila × M. spectabilis] Only cultivated Ornamental
M. × moerlandsii Door. 34 [= M. × purpurea ‘Lemoinei’ × M. toringo] Only cultivated Ornamental
M. × platycarpa Rehder 51, 68, A [= M. domestica × M. coronaria] Eastern North America
M. × purpurea (E. Barbier) 34 [= M. astrosanguinea × M. pumila Only cultivated Ornamental
Rehder ‘Niedzwetzkyana’]
M. × purpurea f. eleyi (Bean) Rehder [= M. ×
purpurea ‘Eleyi’ (M. astrosanguinea × M. pumila
‘Niedzwetzkyana’]
M. floribunda var. lemoinei E. Lemoine [= M. ×
purpurea ‘Lemoinei’]
M. × purpurea f. lemoinei (E. Lemoine) Rehder
[= M. × purpurea ‘Lemoinei’]
M. × purpurea var. aldenhamensis Rehder
[= M. × purpurea ‘Aldenhamensis’]
5
Continued
Apples - Chap 01
6
11/4/03
11:00 am
Page 6
Chromosome
M. × robusta (Carriere) [= M. baccata × M. prunifolia]
J.J. Luby
34 China, cultivated Ornamental, rootstock
Rehder
M. × scheideckeri Spath ex – [= M. floribunda × M. prunifolia] Only cultivated Ornamental
Zabel
M. × soulardii (L.H. Bailey) 34 [= M. ioensis × M. pumila] Central USA, naturalized Fruit, ornamental
Britton and cultivated
M. × sublobata (Dippel) – [= M. prunifolia × M. toringo] Only cultivated Ornamental
Rehder
Taxonomic Classification and History 7
and will commonly graft and plant desirable 1995; Zhou, 1999). Malus × asiatica is proba-
M. sieversii from the forest in their gardens. bly a hybrid complex derived primarily from
Planting desirable trees from root suckers M. sieversii with M. prunifolia and perhaps
may also have been a common practice prior other species.
to, or in addition to, grafting, as M. sieversii Prehistoric remains and historical records,
trees sucker freely. Conversely, people may reviewed by Morgan and Richards (1993),
have cloned and moved some of their horti- provide evidence of the cultivation, dispersal
culturally desirable trees to areas where they and human use of the apple in Asia and
seasonally grazed their animals. These trees Europe over the last several thousand years.
or their open-pollinated descendants may be Archaeological remains of apple that dated
among the horticulturally elite specimens to about 6500 BC were found in Anatolia,
observed in some of the forests today. though it is impossible to know the source of
The passage of trade routes from China to this fruit or whether it was cultivated.
the Middle East and Europe through Central Historical evidence referring to apple culti-
Asia probably facilitated repeated short- and vation dates to the second millennium BC
long-distance dispersal to the east and west, from Anatolia and northern Mesopotamia.
either intentionally or unintentionally, of M. By 500 BC, the apple was probably cultivated
sieversii and its hybrid derivatives. The M. × widely throughout the Persian Empire, as
domestica complex may then have arisen fruit orchards feature prominently in writ-
through hybridization to the east with ings from the period. When Alexander the
species native to China, including Malus Great conquered the Persians around 300 BC,
prunifolia (Willd.) Borkh., Malus baccata (L.) the cultivation of fruits was dispersed
Borkh., Malus mandshurica (Maxim.) Kom. through the Greek world. By this time, the
and Malus sieboldii (Regel) Rehder. To the Greek philosopher, Theophrastus, had dis-
west, hybridization with the local species M. tinguished the sweet cultivated apple from
sylvestris and Malus orientalis Uglitzk. is con- astringent wild forms.
jectured (Ponomarenko, 1983; Morgan and The ascendance of the Roman Empire
Richards, 1993; Hokanson et al., 1997; Juniper spread cultivation of the domesticated apple
et al., 1998). north and west through Europe, where it
During the late 19th and 20th centuries, supplanted and probably hybridized with
M. × domestica cultivars found or bred in the native crab apple, M. sylvestris. Multiple
Europe, Russia, North America, New varieties were recorded by the Roman writer
Zealand, Japan and Australia were intro- Pliny, and they had attained an important
duced throughout the world and form the place in Roman cuisine, medicine and aes-
basis for most current commercial apple pro- thetics by the 1st century AD. The Roman
duction (Way et al., 1990; Janick et al., 1996). goddess Pomona was revered as the deity
Several species are known to have con- associated with apple and other fruits. With
tributed to the M. × domestica complex in the rise and spread of Christianity and Islam
modern breeding programmes including over the next several centuries, apples were
Malus floribunda Siebold ex Van Houtte, carefully maintained, even through wars and
Malus micromalus Makino, Malus × atrosan- difficult times, in the abbey gardens through-
guinea (Spath) C.K. Schneid., M. baccata, out Europe and the orchards of Iberia. These
Malus zumi (Matsum.) Rehder and Malus sar- apparently replaced the native crab apples,
gentii Rehder (Ponomarenko, 1983; Way et al., which had a place in the diet of early Celts,
1990; Janick et al., 1996). Gauls, Franks, Scandinavians and other peo-
In southern and eastern Asia, nai or the ples of northern Europe in fermented, dried
Chinese soft apple, Malus asiatica Nakai, was or cooked forms. Maintenance of fruit gar-
the primary cultivated apple in China and dens was encouraged as a basic monastic
surrounding areas for over 2000 years until skill and many abbeys developed large
M. × domestica was introduced in the late orchards with many M. × domestica cultivars.
19th and early 20th centuries (Morgan and Likewise in the Muslim world of the eastern
Richards, 1993; Zhang et al., 1993; Watkins, Mediterranean and Iberia, fruit growing was
8 J.J. Luby
revered in keeping with Koranic teachings Apples were introduced to Australia, on

and skills of grafting, training and pruning the island of Tasmania and at the present
became highly developed. site of Sydney, in 1788. Orchards were estab-
From the 13th century, apples became lished by settlers in Tasmania and New
more and more widely planted throughout South Wales by the early 1800s. Significant
Europe in gardens of royalty and common- production areas were eventually developed
ers. Raw apples were occasionally con- in Tasmania and the south-eastern main-
sumed, but they were more greatly prized land. In 1814, English missionaries brought
when cooked and sometimes blended with apples from Australia to New Zealand,
spices and sugar or honey. Fermented juice, where two large apple-production districts
or cider, like beer, was preferred to the some- became established in the districts of
times questionable local water-supply. By the Hawke’s Bay and Nelson during the 19th
17th century there were at least 120 cultivars and 20th centuries.
described in western Europe. The rise and Beginning in the 16th and 17th centuries,
spread of Protestantism, which saw the European colonists brought apples to the
apple as the special fruit of God, is credited Americas. Spanish priests introduced them
with expanding apple cultivation across to their missions in Chile and California.
northern and eastern Europe after beginning Spanish and Portuguese settlers introduced
in Germany in the early 17th century. By the apples to their settlements in suitable tem-
end of the 18th century, many hundreds of perate climate zones of South America.
cultivars were recognized throughout European settlers brought apple seeds to
Europe. The Royal Horticultural Society of establish orchards in the eastern USA and
England acknowledged at least 1200 in 1826. Canada. Apples grew well from northern
The 18th and 19th centuries saw apple culti- Georgia to eastern Canada and, as in Europe,
vars recognized and classified based on their were soon highly prized for food and drink
suitability for their end uses (Plate 1.3). and as a source of sugar and alcohol. The
Aromatic dessert apples were more widely first orchards in New England were recorded
appreciated by this time, while good cooking in the 1620s and 1630s and became impor-
types were still appreciated for puddings tant components of the New England farm-
and pastries (Plate 1.4). Flavourful cultivars stead (Plate 1.9). Likewise, they became
with moderate acid and tannin levels were important on the large plantations of the
prized for cider production. The late 19th mid-Atlantic colonies by the mid-1700s,
and early 20th centuries represented the including those of the early US presidents
maximum of diversity in apple cultivation in George Washington and Thomas Jefferson.
Europe, with hundreds of locally popular Jefferson, an astute horticulturist, acquired
cultivars being grown in thousands of small and carefully tested dozens of cultivars for
orchards (Plates 1.5 and 1.6). In the 20th cen- his Monticello gardens in Virginia.
tury, the rise of imported fruit from the In Canada, French colonists established
Americas, New Zealand (Plate 1.7), Australia orchards in the 17th century along the St
and South Africa (Plate 1.8) forced European Lawrence valley. Settlers also established
orchards to increase in size and decrease in orchards around Lake Ontario and in the
number and, to a large extent, to adopt the milder valleys of Nova Scotia and New
very same cultivars that were developed in Brunswick.
and imported from the New World. As settlers moved westward in the USA,
Apples were established in the 1650s near apple orchards were a requirement of home-
Cape Town in South Africa to sustain settlers steading throughout the territories of the
and to supply the ships of the Dutch East Ohio River valley. Jonathan Chapman,
India Company. The commercial apple known as Johnny Appleseed (Fig. 1.1),
orchard district in the Western Cape was devoted his later life, from 1806 to 1847, to
started by Cecil Rhodes and his associates in helping settlers establish thousands of apple
the late 19th and early 20th century to trees on their new farms in the Ohio River
replace a faltering wine industry. drainage. The Great Lakes region of the USA,
nations. Later in the century, the USSR also

became important. By the beginning of the
21st century, China has become the largest
apple producer, with a large proportion of the
crop being exported as concentrated juice
(Plate 1.11). Major southern-hemisphere pro-
duction, much of it for export to northern-
hemisphere countries during their spring and
summer, occurs in South Africa, Chile,
Argentina, New Zealand and Australia.
Production is currently dominated by strains
of just a few cultivars: ‘Delicious’, ‘Golden
Delicious’, ‘McIntosh’ and ‘Jonagold’ devel-
oped in North America; ‘Braeburn’ and ‘Gala’
from New Zealand; ‘Granny Smith’ from
Australia; and ‘Fuji’ from Japan. Though
many other cultivars remain locally impor-
tant, these dominate current production and
are also widely used in breeding programmes
around the world.
From its origins among the millions of
wild M. sieversii trees in the mountains of
central Asia and from the early development
of thousands of local cultivars in Europe and
America, the domesticated apple, as culti-
vated in the 21st century, has shrunk drasti-
cally in diversity.
Fig. 1.1. Only known drawing of Jonathan
Chapman, ’Johnny Appleseed’, supplied by Johnny
Appleseed Heritage Center, Inc.
1.2 Taxonomy and Evolution
1.2.1 Apples in the family Rosaceae

especially the states of New York, Michigan
and Ohio, continues to be a major apple- Apples are members of the genus Malus
production area. Miller, which is placed in the subfamily
In 1847, as settlers moved into the pro- Maloideae of the family Rosaceae (Fig. 1.2).
ductive valleys of western Oregon, Other members of the Maloideae that are culti-
Washington and northern California, vated for their fruit include pears (Pyrus L.
Henderson Llewelling brought 700 trees spp.), quinces (Cydonia oblonga Mill.), loquats
with his family on the Oregon Trail and (Eriobotrya japonica (Thunb.) Mill.), medlars
eventually established the first fruit nursery (Mespilus germanica L.) and species of
in the Pacific Northwest. As irrigation Amelanchier, Aronia, Crataegus and Sorbus. The
schemes were eventually developed, the subfamily Maloideae is one of four in the fam-
Pacific Northwest, especially including the ily Rosaceae. The other subfamilies are
basin of the Columbia River and its tribu- Rosoideae, Spiroideae and Amygdaloideae. The
taries west of the Cascade Mountains and circumscription of these subfamilies has
extending to the Okanagon River valley in defied general agreement among systema-
British Columbia, eventually became one of tists, depending on whether classification
the pre-eminent apple-production areas of schemes emphasize morphological traits,
the world (Plate 1.10). chromosome numbers, intergeneric crossabil-
By the early 20th century, the USA and ity or molecular polymorphisms (Rohrer et al.,
Canada were the two largest apple-producing 1991, 1994; Morgan et al., 1994). The Maloideae
10 J.J. Luby
Family
Rosaceae
Subfamily Subfamily Subfamily Subfamily

Maloideae Rosoideae Spiroideae Amygdaloideae
Genus Genus Genus 20 other

Malus Pyrus Sorbus genera
Section Section Section Section

Malus Sorbomalus Docyniopsis Eriolobus
e.g. M. doumeri e.g. M. trilobata
Series Series Series Series Series Series

Malus Baccatae Sieboldianae Florentinae Kansuenses Yunanenses
e.g. M. sieversii e.g. M. baccata e.g. M. sieboldii e.g. M. florentina e.g. M. kansuensis e.g. M. yunnanensis
Fig. 1.2. Taxonomy of Malus (adapted from Phipps et al., 1991).
are characterized by a hypanthium and al., 1985; Weeden and Lamb, 1987; Dickson et
gynoecium that remain fused to form an in- al., 1991). An allotetraploid origin involving
ferior ovary that develops into a fleshy, inde- ancestral Spiroideae (mostly x = 9) and
hiscent fruit, or pome. Some genera with Amygdaloideae (x = 7) was proposed by Sax
capsules or follicles, however, are apparently (1931, 1933) and is supported by flavonoid
more closely related to genera in the Maloideae chemistry (Challice, 1974; Challice and
than to genera in other subfamilies, based on Kovanda, 1981) and morphological traits
DNA sequence variation (Morgan et al., 1994). (Phipps et al., 1991). DNA sequence variation
The subfamily Maloideae has a high hap- in the rbcL chloroplast gene suggests that
loid base chromosome number of x = 17 and Amygdaloideae and Maloideae are both
is generally considered to be monophyletic advanced groups that arose from x = 9
when morphological traits, chromosome spiraeoid-like ancestors (Morgan et al., 1994).
number (Kalkman, 1988; Phipps et al., 1991) Data from internal transcribed spacer
and DNA sequence variation from the chloro- regions of nuclear ribosomal DNA genes are
plast rbcL gene (Morgan et al., 1994) and S- less comprehensive but support Spiraea as a
RNase (self-incompatibility) gene (Ushijima closer relative to the Maloideae than Rosa or
et al., 1998) are considered. Data from nuclear Prunus (Campbell et al., 1995).
ribosomal DNA sequences, however, support The taxonomic treatment of genera within
a single phylogeny for most of the genera, Maloideae has varied from five cited in
including Malus, but a separate phylogeny Linnaeus’s original treatment up to 33
for the genera Eriobotrya, Rhaphiolepis and (Robertson et al., 1991). Varying morphology
Vauquelinia (sometimes placed in the and numerous instances of intergeneric
Spiroideae) (Campbell et al., 1995). hybridization complicate delimitation of gen-
Based on cytology and analysis of mor- era. Robertson et al. (1991) describe 28 genera,
phological characters, the Maloideae probably including Malus. Species currently included
have a polyploid origin (Phipps et al., 1991). in Malus were included in Pyrus by Linnaeus
Isozyme studies in Malus support an and others until the mid- to late 19th century.
allopolyploid origin, based on the presence Campbell et al. (1995) considered molecular,
of duplicated gene systems, allele segrega- morphological and wood anatomical data in
tions and fixed heterozygosities (Chevreau et determining relationships among genera.
Parsimony analyses of nuclear ribosomal persistent calyces on the fruit, and series
DNA sequence variation placed Malus close Baccatae, containing several Asian species,
to Heteromeles, Chaenomeles, Photinia, Cydonia with fruit consisting of three to five carpels
and Pyrus. A numerical taxonomic treatment and deciduous calyces.
of morphological and wood anatomical stud- 2. Section Sorbomalus, including series
ies placed Malus in a cluster that includes Sieboldianae, with species native to Japan,
Crataegus, Mespilus, Amelanchier, Peraphyllum series Florentinae, with Malus florentina
and Rhaphiolepis. A parsimony analysis, with (Zuccagni) C.K. Schneid. from south-east
both morphological and molecular data Europe, series Kansuenses, containing small-
pooled, placed Malus close to Chaenomeles, fruited Chinese species (and the North
Pyrus and Aria. American Malus fusca (Raf.) C.K. Schneid.),
with deciduous calyces and persistent fruit,
and series Yunnanenses, species from China
1.2.2 Species in the genus Malus with persistent calyces and generally persis-
tent fruit.
The delimitation of species within Malus 3. Section Eriolobus, containing only Malus
has been problematic, with various treat- eriolobus (Poir.) C.K. Schneid. from the east-
ments recognizing from as few as eight to ern Mediterranean.
as many as 78 primary species 4. Section Choromeles, containing exclusively
(Ponomarenko, 1986; Phipps et al., 1990). North American species.
Many hybrid species, derived naturally or 5. Section Docyniopsis, containing the species
artificially, are recognized (Phipps et al., Malus tschonoskii (Maxim.) C.K. Schneid.,
1990; Way et al., 1990). Many of the com- Malus doumeri (Bois) A. Chev., Malus melliana
monly described primary species and (Hand.-Mazz.) Rehder and Malus formosana
hybrid derivatives are listed in Table 1.1. Kawak. & Koidz. of Japan, Taiwan and
The classification and species retained here South-East Asia.
are consistent with the taxonomy database
of the US Department of Agriculture Robertson et al. (1991) revised the genera
Germplasm Resources Information Net- in Maloideae based primarily on a compre-
work (USDA, ARS, National Genetic hensive numerical taxonomic treatment of
Resources Program. Germplasm Resources 115 morphological traits, including foliage,
Information Network (GRIN), 2000) at inflorescence and fruit by Phipps et al. (1991).
http://www.ars-grin.gov on the World In the genus Malus, they retained three sub-
Wide Web. A primary centre of species rich- genera: (i) Malus; (ii) Sorbomalus; and (iii)
ness and diversity is in south-west China, Chloromeles. Several former Malus species are
with several species ranging east to placed in other genera: Eriolobus includes E.
trilobata (Poir.) M. Roem. (= Malus trilobata)
Manchuria and Japan and others extending
and Docyniopsis includes D. tschonoskii
to western Europe. A secondary centre exists
(Wall.) Decne. (= M. tschonoskii) and presum-
in North America, with four native species.
ably would include M. doumeri, M. formosana
The species of Malus have been arranged
and M. melliana. They suggested that further
in varying numbers of sections or subgenera,
work may support inclusion of the genus
some of which are, in turn, divided into
Docyniopsis as part of the genus Docynia and
series (Fig. 1.2). Most recent authors modify
elevation of subgenus Chloromeles to genus.
the treatment of Rehder (1940) and assign
The difficulty in species delimitation in
Malus species to five sections of the genus
Malus arises from the great diversity, poten-
based on morphological traits and flavonoid
tial for hybridization and polyploidy and
similarities (Phipps et al., 1990):
presence of apomixis in the genus (Campbell
1. Section Malus, consisting of series Malus, et al., 1991). These phenomena may be
including many European and Asian species indicative of a fairly recently derived genus
(including M. sieversii and M. × domestica), in which species have developed rapidly
with fruit having five carpels and mostly through adaptive radiation and are primarily
12 J.J. Luby
isolated by geography. Genetic barriers are tica cultivars ‘Golden Delicious’, ‘McIntosh’
not well developed, as putative natural and ‘Delicious’ each had a different haplo-
hybrids are common and artificial interspe- type. One accession of Malus micromalus
cific hybrids are easily produced (Korban, shared its haplotype with two accessions of
1986; Way et al., 1990). Molecular polymor- M. baccata, but the other two accessions of M.
phisms have been used to identify affinity micromalus each had a novel haplotype, one
and phylogeny among taxa in Malus. At the of which was shared with M. floribunda.
molecular level, many relationships are simi- Several researchers have attempted to
lar to those based on traditional classifica- determine relationships among Malus
tions. Nevertheless, some anomalous species or accessions using DNA polymor-
relationships remain problematic, even at the phisms generated by the polymerase chain
DNA level. In many studies, the lack of reso- reaction using either primers with random
lution may be due to limited sampling where sequences (random amplified polymorphic
only one accession is used to represent a DNA, RAPD) or primers that specifically
taxon. In addition, the veracity of a wild amplify DNA in segments containing multi-
species accession may be questionable if it ple repeats of simple base motifs (simple
was obtained as seed from a botanic garden sequence repeat, SSR). Dunemann et al.
or even in native sites occupied by multiple (1994) examined RAPDs in 27 M. × domestica
Malus species or in close proximity to cultivars and in 18 accessions of other
domesticated apple trees. species and found that they supported the
The occurrence of various flavonoids in close relationship of M. × domestica with M.
Malus species (Williams, 1982) was in gen- pumila and M. sylvestris and the distance of
eral agreement with relationships estab- section Malus species from M. ioensis in sec-
lished by morphology except that M. tion Chloromeles and E. trilobata (= M.
florentina showed greater affinity with trilobata). RAPDs observed by Zhou and Li
species in section Docyniopsis than in (2000) and Oraguzie et al. (2001) support the
Sorbomalus, where Rehder (1940) placed it. A close relationship of M. sieversii, M. prunifolia
phylogenetic analysis based on chloroplast and M. sylvestris with M. × domestica and a
DNA restriction-site polymorphisms identi- slightly more distant relationship with M.
fied three lineages (Matsumoto et al., 1997). orientalis and M. baccata. Observing SSR
One included only species in section Malus polymorphisms among 142 species and
of the genus, including M. asiatica, M. hybrid accessions, Hokanson et al. (2000)
baccata, M. mandshurica, M. sargentii, Malus deduced that accessions of M. fusca formed a
prattii, Malus transitoria and Malus distinct group, as did accessions of North
toringoides. A second included M. pumila and American section Chloromeles species, M.
M. prunifolia from section Malus, but also ioensis, Malus coronaria and M. angustifolia.
included M. tschonoskii from section Beyond these groupings, SSR markers were
Docyniopsis. The third group included the not useful in establishing species relation-
species Malus angustifolia (Aiton) Michx. and ships or phylogeny.
Malus ioensis (A.W. Wood) Britton of the The initial analyses of molecular data,
North American section Chloromeles but also described above, provide only minimal addi-
included Malus yunnanensis (Franch.) C.K. tional insights into relationships and phy-
Schneid. and M. florentina of section logeny beyond those suggested by classical
Sorbomalus, and even Eriolobus trilobata (= M. analyses based on morphological traits.
trilobata of section Eriolobus). Although more sophisticated molecular
Restriction-site polymorphisms in mito- analyses may provide more precision in the
chondrial DNA from 14 genotypes yielded future, high resolution may still be quite dif-
11 haplotypes and further illustrated the con- ficult to obtain, especially among the many
fusion in relating molecular data to conven- Eurasian species in section Malus, where
tional classifications (Kato et al., 1993). The divergence may have been relatively recent,
12 accessions from section Malus accounted enforced primarily by isolation or obscured
for nine distinct haplotypes. The M. × domes- by hybridization.
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2 World Production, Trade, Consumption

and Economic Outlook for Apples
Desmond O’Rourke
Belrose Inc., Pullman, Washington, USA
2.1 Introduction 15
2.2 Apple Production Trends 15
2.3 Commercial Practices 17
2.4 Cultivars 18
2.5 Controlled-atmosphere Storage 19
2.6 Market Uses of Apples 19
2.7 The Conundrum of Export Markets 20
2.8 Export Economics 21
2.9 International Trade Laws and Trade Barriers 22
2.10 Global Apple Consumption and Demand 23
2.11 Consolidation in Food Distribution 25
2.12 Stimulating Apple Demand 26
2.13 Need to Balance Supply and Demand 27
2.14 Future Outlook 27
2.15 Concluding Comments 28
2.1 Introduction and ‘Fuji’) and advances in irrigation tech-

nology have permitted apple production to
Apples grow readily throughout temperate expand successfully into warmer climates
climatic zones. However, commercial apple (O’Rourke, 1994).
production is increasingly concentrated in
countries and in growing districts that have
a strong comparative advantage in apple 2.2 Apple Production Trends
production and marketing. Falling trade
barriers have meant that it has become World apple production has been on a
increasingly difficult for less efficient prolong-term growth trend since the Second
ducers to find shelter from more efficient World War. The rate of growth slowed in
external competitors. The development of the 1980s, but leaped ahead in the 1990s,
more heat-tolerant cultivars, the increasing due to just one factor, the phenomenal
popularity of varieties that require a long expansion of production in China. At the
growing season (such as ‘Granny Smith’ beginning of the 1990s, Chinese apple pro-

16 D. O’Rourke
duction was about 4 million t. By the end of surviving producers have intensified their
the decade it had grown more than five- production practices. Production has also
fold. China has provided all the increase in grown in countries like Turkey, Iran, India
acreage of apples harvested around the and Pakistan, where irrigation water is avail-
world since the mid-1980s (Fig. 2.1). In the able and higher-value fruit crops have
1990s, apple acreage harvested in the rest of replaced less profitable field crops.
the world has been declining (O’Rourke, Apple production in China had been
2000). growing steadily since the Second World
Production increases in most countries War and was about equal to that of the USA
have resulted primarily from more intensive in 1990. However, no one predicted that the
production methods, rather than from any rate of growth would be maintained from
significant net increases in planted area. the higher base in 1990 so that China’s pro-
Orchards have become fewer and larger. duction would exceed 20 million t by the end
Producers have concentrated production on of the decade. Over the same period, produc-
large, level blocks on valley floors rather tion of apples in the rest of the world
than persisting with smaller, more difficult changed little (Fig. 2.2). Food and
hillside blocks. Agriculture Organization (FAO) data indi-
Apple production has been static or cate that China’s share of world apple pro-
declining in many countries of both Europe duction has gone from 10.7% in 1990 to
and the former Soviet Union in the last 36.7% in the year 2000.
decade as traditional plantings have been Many factors contributed to this growth.
exposed to competitive realities (FAO, a). It There was a pent-up demand for fresh fruit
has been rising modestly in North America among the huge (1.2 billion) Chinese popula-
over this same period and at a brisk pace in tion. Demand for apples was very respon-
southern hemisphere producing countries. sive to various indicators of economic
Improvements in transportation, relatively growth (Han et al., 1999). Fruit supplies had
weak exchange rates and the demand of been limited in order to provide security of
supermarkets for year-round supplies have grain supplies. Beginning in 1978, with the
given southern hemisphere producers an introduction of the ‘production responsibil-
incentive to expand production. ity’ system for agriculture, the Chinese gov-
However, even in Europe, apple produc- ernment gradually relaxed the restrictions on
tion has continued to grow in major produc- what peasant farmers could grow, how they
ing countries such as France and Italy as could market their product and what price
7000
6000
Area harvested (’000 ha)
5000
4000
3000
2000
1000
0
1967–1969 1977–1979 1987–1989 1997–1999
Fig. 2.1. Apple area harvested in China , rest of the world and total world, 1967–1999 (from FAO, 2000).
Apples: World Production, Trade and Consumption 17
70
60
Production (million t) 50
40
30
20
10
0
1990 1992 1994 1996 1998 2000
Fig. 2.2. Apple production harvested in China , rest of the world and total world, 1990–2000 (from
FAO, 2000).
they could receive. A smallholding that 2.3 Commercial Practices

would provide a meagre living from rice
sold at low government-ordained prices While many small or part-time operators in
could generate a many times greater income many countries continue to produce apples
from fruit sold at market prices. In addition, in traditional ways, the majority of commer-
the leaders of communes and townships cial growers are increasingly exposed to best
encouraged fruit production and processing commercial practices from around the world.
as a way to boost rural incomes. Greater ease of international communica-
Most of China’s apple production occurs tions, individual and group study tours, geo-
in six provinces: Shandong, Shaanxi, Henan, graphical diversification by multinational
Hebei, Liaoning and Shanxi, with about half corporations, international alliances among
occurring in Shandong and Shaanxi alone growers, packers, nurseries, chemical com-
(China State Statistic Bureau, 1999). panies, research and extension workers and
Technically, ownership of the land is distrib- so on, together with international coverage
uted in small lots (less than 0.1 ha) to indi- in most major fruit journals, have all played
vidual peasants. However, titles are a part in increasing the homogeneity of com-
frequently unclear. Through assorted institu- mercial practices among the better growers.
tional arrangements, some large contiguous Importers, wholesalers and retailers that
blocks are under common leadership or buy fruit across international borders are
management. No precise evaluation of the increasingly concerned about how that fruit
calibre of production practices employed has is produced and handled on its way to mar-
been reported, but these appear to vary ket. Standards for integrated production, such
widely. Sophisticated practices appear to be as the International Standards Organization
most widespread in the coastal province of (ISO) 9000 and Hazard Analysis and Critical
Shandong. United States Department of Control Point (HACCP) protocols, are being
Agriculture (USDA) data indicate that ‘Fuji’ demanded by more buyers and more govern-
in 1999 accounted for 45% of acreage and ment monitoring agencies. All these are shin-
production and ‘New Red Star’ for 12%. ing a spotlight on current commercial
Cultivars developed in China, such as practices and forcing the apple industry to
‘Chalajin’, ‘Guoguan’, ‘Qinguan’ and move to generally accepted best commercial
‘Jinguan’, each accounted for less than 10%. practices that improve the sustainability of
‘Gala’ was the western variety most popular production methods and reduce dependence
for new plantings (Rutledge, 2000). on the use of chemical pesticides.
18 D. O’Rourke
2.4 Cultivars the apple business at the expense of existing

cultivars. The battle between cultivars is
International information flow has also had a being fought in the nurseries, in the
profound effect on the popularity of different orchards, in the packing and storage facilities
apple cultivars. In most countries, growers and on the retail shelf. The shake-out among
continue to produce many cultivars that have cultivars can be expected to continue. Some
been traditional in their region. However, of the older, local cultivars are rapidly disap-
newer cultivars have gradually been intro- pearing, while others have proved more
duced for varied production and marketing resilient. In turn, some of the newer cultivars
reasons. For example, ‘Red Delicious’ and have flourished in certain districts while
‘Golden Delicious’ became popular in the some have not.
USA in the 1950s and 1960s, partly because Official data on cultivars are either very
they provided attractive display opportuni- limited or not available at all for many coun-
ties for the booming supermarket retail busi- tries. Estimates of cultivar trends for 34
ness. ‘Granny Smith’ entered the market major producing countries (not including
initially as an off-season cultivar. Production China) were developed by O’Rourke (2001).
expanded in the northern hemisphere in the These suggest that the volume of production
1970s and 1980s as the demand grew for a of ‘Red Delicious’, ‘Golden Delicious’ and
tarter apple. The arrival of ‘Jonagold’, ‘Gala’, ‘Granny Smith’ apples will be stable in the
‘Fuji’ and ‘Braeburn’ in the 1980s and 1990s next few years, but that there will be signifi-
coincided with the growth of hypermarkets cant gains in volume of ‘Gala’, ‘Braeburn’,
and a large expansion of the retail shelf space ‘Pink Lady®’ and, to a lesser extent, ‘Fuji’
devoted to produce. (Table 2.1). If China is included, ‘Fuji’ pro-
Since per capita consumption of apples in duction will continue to leap ahead because
most developed countries is flat, in general China is estimated to have 45% of its produc-
new cultivars can only expand their share of tion in this one cultivar.
Table 2.1. Major producing countries excluding China, cultivar trends

(’000 t) (from O’Rourke, 2001).
2000 2005 2010

Cultivars Estimated Projected Projected
‘Red Delicious’ 5,334 5,422 5,423

‘Golden Delicious’ 4,982 5,222 5,212
‘Granny Smith’ 1,719 1,805 1,828
‘Rome Beauty’ 565 560 540
‘Cox’s Orange’ 218 232 226
‘McIntosh’ 543 655 665
‘Jonathan’ 678 737 727
‘Idared’ 805 1,087 1,148
‘Fuji’ 1,433 1,671 1,857
‘Gala’/‘Royal Gala’ 1,688 2,188 2,594
‘Braeburn’ 429 574 710
‘Elstar’ 413 494 535
‘Gloster’ 210 221 214
‘Jonagold’ 1,039 1,168 1,208
‘Jonagored’ 199 240 285
‘Pink Lady® ’ 61 129 194
All other 5,997 6,726 7,063
Grand total 26,313 29,131 30,429

The outcome of these changes in cultivars able, apples that are not marketable as fresh
is that all the major exporting countries can are used for animal feed or are wasted. In a
offer plentiful supplies of both traditional few countries, such as the USA, Germany
and newer cultivars. Thus, year-round com- and Australia, there has been a large market
petition in almost every cultivar is likely to for apples processed into forms such as
intensify. slices, pie fillings, dried apples, apple sauce,
juice or cider. Specific cultivars were, and
continue to be, grown because of their suit-
2.5 Controlled-atmosphere Storage ability for these processed products.
The development in the 1960s of an effec-
Controlled-atmosphere (CA) storage was tive technique for concentrating apple juice
originally adopted as a tool to permit compa- to a six-to-one ratio has revolutionized the
nies to market selected cultivars of apples for global market for apple juice. Because of its
an additional month or two. As the technol- bulk and low value, single-strength apple
ogy has evolved, CA has enabled more culti- juice had to be marketed near the point of
vars to be held for longer periods. It has been production. Regional processors were effec-
an excellent tool for managing the flow of tively buffered from invasion by outside
product to market. However, the advantages suppliers. However, a turnkey plant to
of CA in lengthening the sales period have process concentrated apple juice (CAJ) could
been offset by the increasing availability of be placed anywhere there was a large supply
new-crop apples from the opposite hemi- of cull apples. The resultant product could
sphere within 6 months of harvest. In recent be shipped in bulk containers around the
years, CA storage has been used increasingly world at low cost. It could be stored without
to ensure better firmness or to control fruit refrigeration and reconstituted on demand.
disorders that affect apples in regular cold Many CAJ plants (most focused primarily on
storage. CA has become a highly complex exports) were erected in major apple-produc-
tool for ensuring that apples meet customer ing countries, such as Argentina, Chile and
quality standards at any time during the mar- Poland. Global supplies of CAJ rose rapidly
keting year. As CA has become more perva- in the 1970s and 1980s (Table 2.2). Growth
sive, buyers have been less willing to pay a has slowed in the 1990s as the apple industry
premium for these quality improvements. in total has shrunk in eastern Europe and the
CA capacity continues to grow in Europe, former Soviet Union.
North America and the southern hemisphere This world pool of CAJ enabled beverage
because larger operators want the flexibility companies and dairies that had formerly
in marketing that CA provides (USDA lacked access to regional supplies of raw pro-
MNS). The volume of apples still in CA stor- cessing apples to build new beverage lines
age in the northern hemisphere during later based on the imported CAJ. They competed
months of the season continues to rise. This directly with the traditional apple processors
has tended to reduce the price advantage that had previously controlled locally avail-
formerly earned by late-season CA fruit. able raw materials. They provided added
That fruit is now beginning to crowd the competition in the market for apple-juice
market and depress the price of new, off-sea- products, but created little new demand for
son apples from the southern hemisphere. domestically produced CAJ because they
Essentially, the seasonal niches that CA stor- could get their supplies more cheaply from
age once made possible have shrunk. the world market. Juice became the primary
use for processed apples, and the world CAJ
price became an important influence on the
2.6 Market Uses of Apples price of apples for other processing uses
(Baumes and Conway, 1985).
The primary market for most apples pro- The percentage of apples processed in
duced around the world is for domestic fresh each country is not known precisely.
use. Where processing facilities are not avail- However, from the USDA Global Agriculture
20 D. O’Rourke
Table 2.2. Concentrated-apple-juice production and ending stocks, selected countries, 1992–2000 (t)
(from USDA FAS).
Country/region 1993/94 1994/95 1995/96 1996/97 1997/98 1998/99 1999/2000p
Production
Germany 60,686 74,571 69,901 68,630 73,903 67,025 70,000
Austria 23,450 14,300 11,500 25,000 30,000 24,700 25,000
Argentina 62,800 65,350 64,400 86,737 59,493 84,800 51,600
USA 146,612 166,547 156,150 134,310 131,966 145,285 150,000
Poland 135,000 108,000 88,000 136,413 145,000 130,000 110,000
N. hemisphere 509,306 509,999 454,797 451,090 519,852 515,128 500,500

S. hemisphere 145,054 141,805 157,040 189,414 152,243 163,274 132,162
World total 654,360 651,804 611,837 640,504 672,095 678,402 632,662
Ending stocks
Germany 164,056 181,286 91,347 81,021 115,000 100,114 87,575
Austria 27,900 25,600 15,900 9,200 13,500 11,200 8,400
Argentina 135 385 0 6,948 0 0 0
USA 0 0 0 0 0 0 0
N. hemisphere 200,531 208,446 107,457 90,241 132,520 129,834 97,495

S. hemisphere 675 885 500 7,348 8,165 3,095 1,595
World total 201,206 209,331 107,957 97,589 140,685 132,929 99,090
p, preliminary.
Information Network (GAIN) data on major For example, Chile and New Zealand
producing countries, we can estimate what have small domestic markets and must find
the general level is by region. In 1998/99, it fresh export outlets for more than half of
ranged from 6% of all apple production their total apple production. In countries
being processed in China to 35% in North such as the USA, France, Italy and South
America and the southern hemisphere and Africa, the domestic market is large but not
to over two-thirds in eastern Europe. In growing fast enough to absorb the increased
North America, about half of all processed supplies at a profit. Certain growing dis-
apples were converted into apple juice. In tricts, such as Washington State in the USA
countries such as China and those in eastern or the Alto Adige in Italy, whose productive
Europe or the southern hemisphere, most capacity far exceeds what the domestic mar-
processed apples were converted into CAJ ket can absorb, must also export fresh apples
and most of that CAJ was then exported. to remain financially viable.
A final major use for apples is for export
fresh. The percentage of world apple pro-
duction entering international trade in fresh 2.7 The Conundrum of Export Markets
form has been between 8 and 10% for many
years (FAO, b). This has occurred because One-fifth of the world’s population lives in
many of the best target markets for exports, developed countries where incomes are high
such as the USA and the European Union, and stable but demand for fresh apples and
have also had an increase in their own apple for apple products is flat. The remaining
production. However, exports have also four-fifths of the world’s population lives in
become critical to the economic vitality of developing countries, where incomes are
the apple industry in many countries and relatively low and unstable, although gener-
growing districts. ally rising. When incomes in the developing
world rise, so does the demand for apples. country, trade will take place. However,
The conundrum for apple marketers is that importing firms may prefer to buy imported
stable markets for fresh apple exports are rather than domestic products for other rea-
not growing and growth markets are not sons, such as timeliness, availability, variety,
stable. For example, economic setbacks in quality, attractive promotion, as an alterna-
Mexico in 1994, in Asia in 1997 and in Russia tive source of supply, because of personal or
and Latin America in 1998 reduced apple business links or for many other less easily
exports to these countries dramatically quantified reasons. In turn, importing firms
while the crises lasted. may prefer a higher-priced domestic product
Eight of the top ten apple-importing for similar reasons. Exporting firms may also
countries in 1998 were developed countries be willing to make uneconomic sales in
(Table 2.3). The exceptions were Russia and order to maintain the loyalty of customers.
Brazil, both of which suffered severe import In produce, the ‘new crop’ designation
declines in subsequent years. In contrast, has always had special appeal. Traders
four of the top ten major apple-exporting believe that new arrivals stimulate fresh
countries in 1998 were developing countries: interest among otherwise jaded consumers.
Chile, South Africa, Argentina and Iran. The Hence, the continuing interest in new-crop
wide differences in costs and sources apples from the southern hemisphere. Other
ensures that global trade channels are sup- factors affect the willingness of exporters to
plied with a wide range of qualities and participate in trade. Governments may pro-
prices of fresh apples. vide direct export subsidies, as in the case of
the European Union. They may provide indi-
rect subsidies, as the USA does through pro-
2.8 Export Economics motional subsidies (the Market Access
Program) or through tax relief for exports
The economics of product exporting is rela- (OECD Committee for Agriculture, 1991).
tively simple in theory. If an exporter can Governments also frequently encourage
deliver goods to a receiving country at a cost exports in subtle ways because they need to
(including shipping, handling and other earn scarce foreign exchange that can be
fees) that is below the price in the importing used to purchase strategic imports.
Table 2.3. Top ten fresh-apple importers and exporters, 1998 (by volume and value) (from O’Rourke, 2000).
Exporters Importers
Volume Value Volume Value
Rank Country (t) (US$ ’000) Country (t) (US$ ’000)
1 France 766,207 488,559 Germany 707,763 438,656

2 USA 582,234 350,454 UK 460,369 390,920
3 Chile 575,601 233,443 Russian Fed. 358,758 137,892
4 Italy 540,138 258,773 Belg-Lux 248,411 204,954
5 The Netherlands 338,901 186,582 The Netherlands 235,922 163,745
6 Belg-Lux 335,470 238,857 Chinaa 158,812 113,032
7 New Zealand 291,720 204,083 USA 141,971 95,390
8 South Africa 242,000 124,470 Spain 132,909 79,463
9 Argentina 227,520 118,093 Brazil 126,186 55,433
10 Iran 190,000 30,000 Canada 115,278 84,749
Top ten 4,089,791 2,233,314 Top ten 2,686,379 1,764,234

Total world 5,176,391 2,660,958 Total world 4,506,625 2,807,216
Top ten % 79.0 83.9 Top ten % 59.6 62.8
aIncludes the province of Taiwan.
Belg-Lux, Belgium and Luxemburg.
22 D. O’Rourke
Another major influence on trade is fluc- consumers to lose, producers to become less
tuation in relative exchange rates. Such fluc- efficient and trading partners to retaliate in a
tuations change the price signals that buyers destructive spiral. The General Agreement
and sellers receive from the market. For on Tariffs and Trade (GATT) was set up in
example, in 1990, one US dollar equalled 2.6 1947 by a small core of developed countries.
South African rands and 337 Chilean pesos. They had suffered a particularly severe bout
By the end of the 2000 marketing season, one of such ‘beggar-my-neighbour’ policies in
US dollar equalled approximately 7 rands response to the Great Depression. GATT
and 560 pesos. If a US importer paid US$10 sought to develop rules governing interna-
per box for apples in both 1990 and 2000, the tional trade and to gradually phase out pro-
South African exporter would have received tectionist devices. In a series of negotiating
almost three times as many rands in 2000 as rounds over the next five decades, GATT
in 1990 but the Chilean exporter only 66% was successful in reducing many trade barri-
more pesos. The US importer would have ers and in recruiting most market economies
received a signal that market demand was as members. At the core of GATT’s success
flat, the Chilean exporter a signal that was the most favoured nation (MFN) princi-
demand was growing modestly and the ple. Members agreed to grant all other mem-
South African exporter a signal that demand bers the same trade concessions (in terms of
was soaring. access, quotas, tariffs, etc.) as they granted to
A stronger currency gives an importing their most favoured trading partner.
country an advantage in bidding for prod- GATT, however, made little progress in
ucts on the world market. Conversely, a liberalizing trade in agricultural products
weaker currency gives an exporting country until the Uruguay Round was completed in
a short-term advantage in undercutting com- 1994 and, even then, made only a modest
peting suppliers. The advantage is usually start. The agency into which the GATT was
short-term because domestic inflation converted, the World Trade Organization
rapidly offsets any nominal increase in (WTO), has failed so far to build on the small
import price. In contrast, a stronger currency gains of the Uruguay Round. One critical
makes life difficult for exporting firms in that exemption to MFN that was allowed by
country, while a weaker currency penalizes GATT has had a lasting impact on the liber-
importing firms. Market signals that are dis- alization process in agricultural products.
torted by currency fluctuations are particu- GATT allowed member countries to form
larly troublesome in perennial crops such as regional free-trade agreements, which could
apples. While producers can respond rapidly liberalize more rapidly than the general
to an apparent increase in price by increasing GATT level. However, just two of those free-
plantings, the additional production gener- trade areas, the European Union and the
ated may overhang the market for many North American Free Trade Agreement
years thereafter. Some of the recent increase (NAFTA), now account for half of world
in world apple production has resulted from trade. The European Union, in particular, has
such distorted price signals. used its strong influence within GATT–WTO
to protect its internal agricultural market by
impeding global trade liberalization. It has
2.9 International Trade Laws and Trade aggressively pursued many new bilateral
Barriers trade links with third countries because
these enable it to exempt sensitive agricul-
In most countries, putting foreign suppliers tural products. The European Union has
at an economic disadvantage relative to found powerful allies in countries like Japan,
domestic suppliers is seen both as fair and as South Korea, Taiwan and even China, which
good politics. Thus, governments are willing fear that their agricultural industries could
to use trade barriers to help domestic pro- not withstand global competition.
ducers. However, experience over two cen- Modest liberalization in agricultural
turies has shown that such policies cause trade has accompanied the implementation
of the Uruguay Round of GATT and of the available for most countries. The USDA and
various regional free-trade agreements. The the FAO both use a balance-sheet approach
European Union has lowered its tariff barri- to estimate consumption indirectly. Total
ers for non-member countries but has left its supplies are considered to equal beginning
variable levies intact. Other countries have inventory plus production plus imports in
reduced their tariffs on agricultural products the period. Fresh consumption is assumed to
as part of their GATT–WTO commitments. be the balance of those supplies remaining
Regional free-trade areas, such as NAFTA after exports, processing use, withdrawals,
and Mercosur (which includes Argentina, waste and losses and closing inventory are
Brazil, Paraguay and Uruguay), have subtracted. For most fresh products, begin-
reduced barriers to member countries. For ning and ending inventory are assumed to
example, Mexico’s duties and tariff-rate be zero, that is, there is no carry-over from
quotas on fresh apples from its NAFTA part- one year to the next. Since consumption is
ners, the USA and Canada, will be phased measured as a residual, any errors in mea-
out by 2004. However, countries that are not surement in any of the other variables affects
members of NAFTA will find themselves at the estimate of consumption.
an increasing disadvantage in selling to the FAO data report per capita consumption
Mexican market. Many countries have of all apples, but do not separate fresh from
moved to resolve this problem by negotiat- processed. From USDA GAIN reports, we
ing bilateral agreements with Mexico. Other estimated per capita apple-consumption
regional free-trade agreements, such as trends between 1990 and 1999 for 32 apple-
Mercosur, will also put non-members at an producing countries (Table 2.4). For 20 of the
increasing disadvantage. Even bilateral 32 countries, the trend was downwards. The
agreements discriminate against those coun- major increases were in China (up 400%),
tries that are not included. The longer a Brazil (up 40%), Taiwan (up over 70%) and
comprehensive global solution to agricul- Turkey (up about 20%). All but Taiwan were
tural liberalization is delayed, the more diffi- low-income, developing countries. We esti-
cult it will be to unravel these growing trade mated comparable data for 22 countries that
distortions. import almost all of their apple supplies
(Table 2.5). In general, they showed a posi-
tive trend for the period 1990–1996.
2.10 Global Apple Consumption and Thereafter, most suffered consumption
Demand declines related to the widespread economic
setbacks of the late 1990s.
We distinguish here between apple con- Demand is a broader concept than con-
sumption and apple demand because they sumption. It describes the relationship
have quite different implications for the between the quantity consumed and the
global apple industry. Consumption is a sta- price paid. Demand is also a much more dif-
tic, one-dimensional measure of the volume ficult concept to measure because it requires
of apples consumed in any time period. Per accurate data series on consumption, prices,
capita consumption is an average figure income, tastes and preferences and other fac-
derived when the total volume consumed is tors that might shift demand. Clearly, it
divided by the total population. Per capita would be more beneficial to the apple indus-
consumption data are useful indicators for try if the average US consumer were willing
comparing the popularity of apples relative to buy 40 lb. of fresh apples annually at US$1
to other fruits, in different time periods or in per lb. rather than the 20 lb. of fresh apples
different countries. actually consumed. However, in a normal
However, measuring consumption of a demand relationship (what economists call
minor food product like apples is difficult the demand curve), consumers will only be
and tends to have a low priority with most willing to consume more if the price is lower.
government statistical agencies. Thus, official One indicator of the strength of demand is
series of per capita consumption data are not what percentage decrease in price would be
24 D. O’Rourke
Table 2.4. Apple-producing countries: estimated per capita consumption, 1990–1999 (kg per capita)
(from O’Rourke, 2000).
1990 1991 1992 1993 1994 1995 1996 1997 1998 1999p
Austria 35.89 30.97 29.15 36.85 32.38 34.14 23.30 25.00 23.73 21.30
Belgium 23.68 20.33 25.36 25.56 29.51 27.69 23.10 21.10 19.63 20.46
Denmark 17.90 14.75 16.34 15.42 16.33 16.51 17.76 20.89 20.82 21.41
France 17.99 13.61 13.80 14.54 16.47 15.69 16.77 15.26 15.61 15.93
Germany 25.01 18.80 32.45 20.05 22.84 18.93 21.06 17.54 21.03 19.99
Greece 16.90 16.22 21.52 21.30 21.40 21.90 18.60 22.08 23.99 24.87
Italy 22.71 20.26 23.46 19.87 21.48 18.78 21.88 21.77 24.23 25.43
Netherlands 23.09 12.00 34.43 23.88 27.26 24.67 24.10 21.26 19.06 19.04
Spain 17.56 14.76 19.54 18.69 17.42 17.64 17.00 17.01 16.85 16.94
Sweden 18.25 15.53 16.78 16.13 17.20 16.37 15.63 17.61 16.40 16.97
UK 12.18 10.89 11.61 11.19 10.73 9.74 9.73 8.86 10.02 10.04
EU-11 20.12 16.22 21.87 18.07 19.28 17.55 18.15 16.97 18.27 18.31
Norway 15.70 12.80 16.67 16.62 15.59 15.54 14.86 13.87 14.40 14.34
Bulgaria 11.80 8.02 6.70 6.50 5.29 5.95 9.05 9.52 7.62 9.05
Hungary 26.58 24.17 20.36 28.99 15.82 12.81 16.16 16.20 15.93 15.77
Poland n/a n/a 12.85 14.57 9.20 8.84 17.66 13.75 8.55 11.59
Romania n/a n/a 18.30 37.00 17.50 16.40 15.45 18.01 13.61 16.37
Slovakia n/a n/a n/a 12.22 10.09 6.05 12.79 12.97 13.87 14.77
Yugoslavia n/a 11.05 22.01 14.61 10.50 9.50 12.81 11.40 8.99 8.95
Other Europe n/a n/a n/a 20.65 11.92 10.85 15.34 14.32 10.95 12.89
Argentina 7.12 7.28 7.74 9.26 8.22 12.40 11.58 9.09 10.02 9.51
Australia 9.39 9.96 10.25 8.94 9.50 8.25 9.29 8.38 8.44 8.89
Brazil 2.90 2.37 3.03 3.16 4.26 4.35 4.15 4.14 3.91 4.11
Chile 6.83 5.23 6.25 6.52 6.57 6.47 6.44 6.14 6.59 6.49
New Zealand 16.14 13.67 15.12 15.31 15.23 14.44 14.57 14.43 30.59 17.32
South Africa 4.94 4.49 5.33 5.55 5.11 5.94 5.75 5.07 4.64 4.85
S. hemisphere 4.56 4.13 4.78 5.01 5.50 6.14 5.93 5.44 5.59 5.52
China n/a 3.73 5.26 7.23 8.80 10.91 13.13 13.14 14.62 16.05
Japan 6.52 3.54 6.61 6.62 6.51 6.38 6.05 6.04 5.59 5.99
Taiwan 4.37 5.54 6.14 5.52 6.64 5.63 6.52 6.76 7.49 7.48
Turkey 30.81 29.98 33.66 33.33 32.18 31.62 32.76 37.78 35.87 36.48
Asia 2.29 5.02 6.82 8.38 9.29 10.98 12.85 13.50 14.67 15.96
Canada 11.64 10.87 13.02 12.20 12.22 12.52 11.95 11.58 12.01 11.76
Mexico 3.57 4.35 5.18 6.67 5.37 4.62 4.87 6.69 4.36 5.41
USA 9.00 8.32 8.79 8.74 8.91 8.57 8.77 8.40 8.72 8.83
N. America 7.96 7.60 8.27 8.51 8.33 8.34 8.08 8.23 7.91 8.21
Russia n/a n/a n/a 4.86 5.03 4.79 5.72 5.27 4.37 4.11
All 32 countries n/a n/a n/a 9.61 9.95 11.37 11.91 11.95 12.53 13.29
n/a, not available; p, preliminary; EU, European Union.

Table 2.5. Per capita disappearance of fresh apples, selected importing countries, 1990 and 1994–1998
(kg per capita) (from O’Rourke, 2000).
Country and region 1990 1994 1995 1996 1997 1998
East Asia
Singaporea 9.65 12.12 12.36 10.00 10.48 9.39
Hong Konga 9.84 11.85 11.35 9.20 8.59 8.62
Taiwan 3.82 5.85 5.18 6.14 5.46 6.80
Malaysia 1.28 2.45 2.71 2.57 3.09 2.02
Thailand 0.42 0.98 1.09 1.08 0.63 0.61
Indonesia 0.01 0.17 0.23 0.19 0.36 0.10
Subtotal 0.71 1.21 1.26 1.21 1.22 1.08
Middle East
United Arab Emiratesa 13.32 20.40 24.39 38.88 18.20 16.47
Bahrain 8.11 14.15 13.86 13.53 11.93 10.17
Oman 4.41 5.33 3.50 1.35 2.19 2.30
Kuwait 5.20 12.75 12.48 14.04 13.75 11.13
Saudi Arabia 8.27 6.89 6.85 7.00 6.76 5.60
Libya 4.02 2.49 1.48 3.75 1.73 2.06
Egypt 0.00 0.35 0.33 0.31 0.45 0.59
Subtotal 2.52 2.87 2.84 3.11 2.55 2.43
Latin America
Panama 1.93 2.12 2.09 2.12 2.06 2.27
Costa Rica 1.60 2.06 2.04 1.92 1.99 2.46
Colombia 0.75 1.30 1.48 1.43 1.61 1.42
Venezuela 0.59 0.95 1.11 0.66 1.32 1.90
Peru 0.11 0.73 0.63 0.51 0.51 1.63
Dominican Republic 0.14 0.51 0.49 0.73 0.51 0.79
El Salvador 0.95 0.48 0.99 0.69 0.88 0.25
Honduras 0.42 0.25 0.32 0.38 0.67 0.17
Nicaragua 0.06 0.14 0.16 0.15 0.20 0.20
Subtotal 0.56 0.96 1.05 0.92 1.12 1.39
aRe-exports subtracted from imports.
needed to achieve a given percentage most systematic approach to studying these

increase in consumption. At the retail level, a changes has occurred at the Retail Food
10% decline in the price of fresh apples Industry Center at the University of
would be needed to boost consumption by Minnesota in St Paul, Minnesota (see, for
10%. However, because marketing margins example, Larson, 1997; Kinsey, 1998). Three
in food distribution are relatively fixed, a related phenomena are driving these
20% or greater decline in grower price would changes. First is the entry of non-traditional
be needed to boost consumption by 10%. retailers, such as discount chains, hypermar-
kets, supercentres and warehouse club
stores, into food retailing. While their appeal
2.11 Consolidation in Food Distribution to consumers goes beyond price factors,
these formats have used economies of scale
Changes in the food-distribution system are in purchasing and internal management effi-
changing the way in which consumer needs ciencies to sell comparable food products at
are being passed back to producers. The prices below those of conventional super-
26 D. O’Rourke
markets. Many have introduced their new The most obvious demand shifter is growth
concepts internationally. Traditional super- in per capita income. However, in developed
market chains have scrambled to regain their countries, where per capita income and per
competitive edge by getting bigger through capita consumption of fresh apples are
mergers and acquisitions and by seeking already high, income increases have little
similar purchasing and logistical efficiencies. positive effect on per capita consumption. In
Many of them, too, have gone international. contrast, in developing countries, where per
As a result, increasing concentration, intense capita income is low, increases in income do
competition among retail formats and down- lead to substantial increases in per capita
ward pressure on supplier prices have consumption of items like fresh apples.
become the norm all over the world. In the These findings suggest that the major
case of apple pricing, as retailers have got apple-exporting countries would get greater
bigger they appear to have acquired some returns by targeting more of their sales
market power (P.M. Patterson, 2000, unpub- efforts and promotional dollars in develop-
lished data). ing-country markets. In developed-country
In most countries, four retailing organiza- markets, where the income effect is weak,
tions now account for 50% or more of all food sales and promotional efforts need to be tar-
sales. Consolidation has led to ever fewer geted at changing consumers’ tastes and
and larger buyers. Most larger buyers want preferences for apples. Many in the industry
to deal with fewer, larger suppliers that can believe that health claims can be exploited to
meet their product needs for 12 months of the achieve this. However, many other fruits and
year, can invest in sophisticated information vegetables can make comparable health
systems and can provide the many war- claims. In addition, enjoyment is a key rea-
ranties and services that retailers and their son for consuming any food. There is a dan-
customers now demand. This has resulted in ger that too heavy an emphasis on the health
a frantic scramble among suppliers to posi- aspect could create a negative image in the
tion themselves to capture some of the busi- minds of many consumers. Work by Baker
ness of the larger retailers. Consolidation is (1999), suggests that, because consumers
now taking place at every level of the supply vary in the depth of their concern for food
system, including packers, shippers, mar- safety and health, such promotions would
keters, brokers, exporters and importers. In have to be targeted more precisely than they
their drive to become larger and more effi- have been in the past. While the appropriate
cient, the surviving suppliers are becoming promotional themes and strategies remain to
less tolerant of individual growers or grow- be determined, there is no doubt that a major
ing districts that cannot meet ever-higher commitment of funds would be needed to
retailer standards. There is no sign that these change consumers’ attitudes and purchasing
forces for change in the food-distribution sys- behaviour substantially.
tem will abate any time soon. A number of leaders in the global apple
industry have become convinced that they
can gain a competitive advantage for their
2.12 Stimulating Apple Demand company by segmenting their marketing
effort by cultivar. They are forming so-called
While efforts to increase per capita consump- ‘marketing clubs’, in which selected growers,
tion of fresh apples may seem desirable, they marketers and nurseries form an interna-
may be counter-productive if they can only tional alliance to manage the licensing, pro-
be secured by sharp reductions in grower duction and marketing of a specific cultivar
price and a decline in grower revenues. throughout the world. The prototype effort is
What is needed is a positive shift in the the International Pink Lady® Alliance, which
whole demand curve, either persuading con- charges participants membership, manage-
sumers to purchase the same quantity but at ment and promotional fees and regulates
a higher price, or a greater quantity at the quality, packaging and other common stan-
same price, or some combination of the two. dards. The sponsors hope that this effort will
boost demand for the ‘Pink Lady®’ cultivar has increased in response to past favourable
to the extent that the added revenue gener- prices, government supports and exchange-
ated will more than compensate for the rate illusions. That supply will continue to
added cost of belonging to the alliance. increase as more trees reach full bearing
Clubs for other cultivars are being formed. unless specific actions are taken to remove
A number of questions have still to be areas from production. Demand, on the
answered regarding these clubs. Can they other hand, has been static in the developed
generate sufficient added revenue to offset world and has been halted, at least tem-
the added costs? Even if there is a profitable porarily, in the developing world by eco-
market segment for ‘Pink Lady®’, how many nomic setbacks and by the failure of
market segments for other varieties can be agricultural trade liberalization.
exploited profitably? Even if these market
segments can be exploited successfully, will
it be at the expense of other cultivars, so that 2.14 Future Outlook
total apple demand is not improved? Will
promotions for individual cultivars be most The consolidation of the apple industry into
effective as a complement to or as a replace- larger, more viable units at the growing, pack-
ment for current generic apple-advertising ing, storage and marketing levels is likely to
programmes? Research is needed into these continue. Less and less of the apple volume
questions before industry groups invest too will be in the hands of marginal growers,
heavily in the marketing-club approach. marginal producing districts and marginal
producing countries. The surviving producers
will employ every available technology to
2.13 Need to Balance Supply and increase yields per hectare and reduce unit
Demand costs. Experimentation with new cultivars
and new marketing models will continue.
Individual markets are like pools of sea However, the biggest single influence on
water on a rocky beach. At low tide, they can the global apple industry in the next decade
be isolated and function independently. But will be what happens to the 2.3 million ha of
an errant wave can at any time rush in and apple orchards in China. Because many of its
sweep over a number of nearby pools and, at trees are not yet at full bearing, China’s aver-
high tide, all pools may be submerged and age yield per hectare in the year 2000 was still
connected. Markets experience the same sort below 10 t. With normal maturation of
of ebb and flow. Local markets can have tem- orchards, average yields can be expected to
porary gluts or shortages that only affect increase by at least 50% in the next decade.
local prices. However, if those gluts or short- Thus, by 2010, Chinese apple production
ages persist, product will flow from other could exceed 35 million t. This would increase
areas of surplus to areas of shortage. Modern global supplies by an average of 1.3 million t
storage, transportation and communication (about 2.0%) each year for the next decade. In
technology has enabled apple suppliers in the same period, global supplies of many
any part of the world to respond rapidly to other fruits are also expected to increase.
any temporary imbalances elsewhere. If, as expected, China is accepted as a
The reality of globalization of markets is member of the WTO, it will become tied
that most local markets are connected with more closely to the world economy and will
the global market most of the time. Thus, the be less free to pursue the ‘China first’ poli-
global balance of supply and demand has cies of the past. Thus, exports of Chinese
become a powerful influence in all but the fresh apples and of CAJ are likely to increase.
most isolated local markets. Increasingly, This will heighten competition in apple mar-
since 1990, there has been an excess of global kets around the world and will intensify
supply over global demand that has pressure on other producers and producing
depressed prices. The reasons for this imbal- districts to rationalize their operations in
ance have already been explained. Supply order to survive.
28 D. O’Rourke
Clearly, too, any effort to control global action. The Southern Hemisphere Asso-
apple supply or boost global apple demand ciation of Fresh Fruit Exporters (SHAFFE),
will have to include China. If the rest of the at its October 1999 meeting in Atlanta, set
world bears these costs, it will rapidly up a task force to study future joint actions
become clear to them that China is getting (Dall, 1999). In October 2000, a broader
some of the benefits of improved prices organization consisting of representatives
without paying any of the costs (the ‘free from leading European and southern hemi-
rider’ problem). However, China does not sphere apple producers set up the World
yet have the organizational structure capable Apple and Pear Association (WAPA) to
of working with the promotional agencies of tackle the same issues that had become even
other major exporting countries. more pressing in the previous year (Anon.,
2000). WAPA hopes to attract participation
from the USA, China and other leading pro-
2.15 Concluding Comments ducing countries.
The producing districts, industry organi-
The global apple industry is entering zations and firms that survive in the 21st
uncharted waters. It needs new maps, new century will be those that best understand
vessels and a new sailing plan to effectively the changing environment in which they
navigate those waters. It needs general must both compete and cooperate. They will
acceptance of the present realities, a vision be those that continuously strive to lower
of how to move forward and a willingness unit costs, produce higher-quality products
to take bold actions to put that vision into and provide the distribution system and con-
effect. A number of global institutions have sumers with continually improving cus-
sought to build such a vision and plan of tomer service.
References
Anon. (2000) World top fruit association to tackle oversupply. Fruit and Vegetable Markets, Agra Europe
122, 19.
Baker, G.A. (1999) Consumer preferences for food safety attributes in fresh apples: market segments, con-
sumer characteristics and marketing opportunities. Journal of Agriculture and Resource Economics
24(1), 80–97.
Baumes, H.S., Jr and Conway, R.K. (1985) An Econometric Model of the US Apple Market. ERS Staff Report
No. AGES 850110, National Economics Division, ERS, USDA, Washington, DC, 30 pp.
China State Statistic Bureau (1999) China Agricultural Yearbook. China State Statistic Bureau, Beijing,
910 pp.
Dall, P. (1999) Deciduous fruit problems global. Deciduous Fruit Grower 49(2), 6–12.
FAO (Food and Agriculture Organization of the United Nations) (2000) FAOSTAT home page [online].
Available: http://apps.fao.org/
FAO, a. FAO Production Yearbook. Food and Agriculture Organization of the United Nations, Rome.
FAO, b. FAO Trade Yearbook. Food and Agriculture Organization of the United Nations, Rome.
Han, T., Wahl, T.I. and Mittelhammer, R.C. (1999) China’s rural household purchasing decisions for high-
valued food products: implications for the US food industry. In: China’s Role in World Food Markets:
Proceedings of WCC-101 Conference, 3–4 February 1999. IMPACT Center, Washington State University,
Pullman, Washington, pp. 219–232.
Kinsey, J.D. (1998) Concentration of Ownership in Food Retailing: a Review of the Evidence about Consumer
Impact. Working Paper 98(4), Retail Food Industry Center, University of Minnesota, St Paul,
Minnesota, 30 pp.
Larson, R.B. (1997) Key Developments in the Food Distribution System. Working Paper 97(8), Retail Food
Industry Center, University of Minnesota, St Paul, Minnesota, 33 pp.
OECD (Organization for Economic Cooperation and Development) Committee for Agriculture (1991) The
Apple Market in OECD Countries. Organization for Economic Cooperation and Development, Paris,
173 pp.
O’Rourke, A.D. (1994) The World Apple Market. Haworth Press, New York, 237 pp.
O’Rourke, D. (2000) The World Apple Review. Belrose, Pullman, Washington, 116 pp.
O’Rourke, D. (2001) World apples to 2010. World Apple Report 8(1), 5–9.
Rutledge, J.D. (2000) China, Peoples Republic of, Fresh Deciduous Fruit, Annual, 2000. Global Agriculture
Information Network (GAIN) Report CH0618, Foreign Agricultural Service, USDA, Washington,
DC, 34 pp.
USDA FAS. World Horticultural Trade and US Export Opportunities. Circular series: FHORT (various
issues).
3 Genetic Improvement of Apple: Breeding,

Markers, Mapping and Biotechnology
Susan K. Brown and Kevin E. Maloney

Department of Horticultural Sciences, Cornell University, New York State Agricultural
Experiment Station, Geneva, New York, USA
3.1 Traditional Breeding 32

3.1.1 Germplasm 33
3.2 Apple-breeding Programmes: Their Objectives and Introductions 33
3.2.1 USA 34
3.2.2 Canada 35
3.2.3 Australia 36
3.2.4 Belarus 36
3.2.5 Belgium 36
3.2.6 Brazil 36
3.2.7 Bulgaria 36
3.2.8 China 37
3.2.9 Czech Republic 37
3.2.10 England 37
3.2.11 Finland 37
3.2.12 France 38
3.2.13 Germany 38
3.2.14 Greece 38
3.2.15 Hungary 38
3.2.16 India 39
3.2.17 Italy 39
3.2.18 Japan 39
3.2.19 Korea (South) 40
3.2.20 Latvia 40
3.2.21 Lithuania 41
3.2.22 Mexico 41
3.2.23 Netherlands (The) 41
3.2.24 New Zealand 41
3.2.25 Norway 41
3.2.26 Poland 41
3.2.27 Romania 41
3.2.28 Russia 42
3.2.29 South Africa (Republic of) 42
3.2.30 Spain 42
3.2.31 Sweden 42
3.2.32 Switzerland 42
3.2.33 Yugoslavia 43
32 S.K. Brown and K.E. Maloney
3.3 Natural and Induced Variation: Mutations and Sports 43

3.4 Genetics and Inheritance of Traits 44
3.4.1 Heritability studies 44
3.5 Disease Resistance: Present Status and Future Directions 44
3.5.1 Apple scab (V. inaequalis (Cooke) Wint.) 44
3.5.2 Powdery mildew (P. leucotricha (Ell. & Ev.) Salm.) 45
3.5.3 Fire blight (Erwinia amylovora (Burrill) Winslow) 45
3.6 Other Traits 46
3.6.1 Quality 46
3.6.2 Nutrition 47
3.6.3 Reduction of flesh browning 47
3.6.4 Allergenicity 47
3.7 Markers and Genetics 47
3.7.1 Isozymes 47
3.7.2 Scab resistance 47
3.7.3 Powdery mildew 47
3.7.4 Insect resistance 48
3.7.5 Incompatibility alleles 48
3.7.6 Other traits 48
3.7.7 Microsatellites/simple sequence repeats (SSRs) 48
3.7.8 Molecular maps 48
3.7.9 Comparative mapping 49
3.7.10 Bacterial artificial chromosome (BAC) library of apple 49
3.8 Biotechnology 49
3.8.1 Somaclonal variation 49
3.8.2 Regeneration and transformation 50
3.8.3 Transgenes introduced 50
3.8.4 Rootstocks transformed 51
3.8.5 Transgene silencing 51
3.8.6 Challenges 51
3.9 Future Prospects 52
3.9.1 Malus genes being investigated 52
3.9.2 1-Aminocyclopropane-1-carboxylate (ACC) synthase and ACC oxidase 52
3.9.3 Additional genes involved in flowering 52
3.9.4 Summary 52
3.1 Traditional Breeding reviewed in Brown (1975). Chapters on spe-

cific areas of breeding, such as pollen collec-
The genetic improvement of scion cultivars tion or mutation breeding, are reviewed in
is the primary focus of this chapter, but the Janick and Moore (1983).
approaches used are also relevant to root- Apple has several constraints to rapid
stock and ornamental Malus breeding. genetic improvement: a long juvenile period,
Rather than emphasizing breeding method- large tree size, self-incompatibility and
ology, an overview of the current status of inbreeding depression. The last two charac-
scion breeding will be detailed (Plate 3.1). teristics mean that apple breeders cannot use
The review of apple breeding by Janick et al. standard backcrossing to incorporate a major
(1996) provides information on apple origin gene from another cultivar or Malus species
and early development, Malus species, mod- and still maintain cultivar identity. A modi-
ern breeding objectives, breeding techniques, fied backcross is used instead, with a differ-
resistance breeding and biotechnology. ent recurrent parent required in each
Information on apple breeding is also generation of backcrossing.
Genetic Improvement of Apple 33
Self-incompatibility enforces outbreeding are active in the development and use of

and results in heterozygosity (Plate 3.2). triploids in breeding.
While some inbreeding for genetic studies Interspecific hybridization has played a
has been successful (Karnatz, 1994), inbred major role in genetic improvement (Korban,
lines cannot be produced easily. Haploid 1986) and continues to be important in culti-
lines are being developed by microspore cul- var development. Interspecific hybridization
ture, anther culture and parthenogenesis in will be aided by molecular markers that
situ (Lespinasse et al., 1999; Keulemans and allow us to follow introgression of genes
De Witte, 2000). These haploids are being from wild species and enable us to differenti-
doubled to obtain homozygous lines for ate hybrid offspring in crosses where
genetic studies. apomixis or parthenocarpy is possible.
In the past, most important commercial
cultivars arose as chance seedlings, and such
seedlings continue to be commercialized 3.1.1 Germplasm
today – for example, ‘Ginger Gold’ and
‘Cameo’. However, increasingly, popular Way et al. (1990) provided information on
new cultivars, such as ‘Fuji’, ‘Gala’, apple germplasm issues, research, important
‘Jonagold’, ‘Elstar’ and ‘Honeycrisp’, are diseases, pests and horticultural problems
derived from breeding programmes. To and listed major apple germplasm centres.
study genetic diversity, Noiton and Alspach The International Plant Genetic Resources
(1996) analysed pedigrees of 50 apple culti- Institute (IPGRI) has a working group on
vars being used in breeding. ‘Cox’s Orange Malus and Pyrus, with reports available
Pippin’, ‘Golden Delicious’, ‘Jonathan’ and online, suggestions for minimum descriptor
‘McIntosh’ were the most frequent progeni- lists for Malus and links to other germplasm
tors. Few of the cultivars sampled were sub- centres at http://www.cgiar.org/
stantially inbred, but continued inbreeding The Plant Genetic Resources Unit in
was a concern. Breeders are currently work- Geneva, New York, part of the US Department
ing with reduced genetic diversity and must of Agriculture (USDA) Agricultural Research
attempt to expand the genetic base. Service (ARS), maintains 3739 accessions of
While cultivated apples are functionally Malus. Information on accessions may be
diploid (2x = 34), many successful cultivars searched in the Germplasm Resources
are triploids (3x = 51). Triploids occur spon- Information Network (GRIN). A core collec-
taneously from unreduced gametes in 2x-by- tion (208 clones) and 40% of the base collection
2x crosses. Triploids have much larger fruit have been characterized for 25 morphological
and are of value to the industry. Breeders descriptors. Information on the USDA Malus
have tried to increase the likelihood of collection can be searched at http://www.ars-
triploids by crossing tetraploids with grin.gov/npgs/ The collection and manage-
diploids, with varying degrees of success. ment of wild Malus germplasm from its centre
Some combinations produce no triploids, of diversity has provided additional sources of
while others may yield 50–80% triploids. genetic variation for use in breeding
Bartish et al. (1999) used the mutagen oryza- (Hokanson et al., 1997).
lin to induce chromosome doubling in
somatic tissues of ‘Florina’ and ‘Falstaff ’ to
produce tetraploid clones for use in breed- 3.2 Apple-breeding Programmes: Their
ing. The quality of some triploid cultivars Objectives and Introductions
has prompted their use as parents, with the
understanding that many of the seedlings There are a large number of public and private
produced will be aneuploids. Triploid culti- breeding programmes around the world. The
vars are best used as pollen parents because Pacific Northwest Fruit Tester’s Association
the 2x gametes will have a selective advan- (PNWFTA) summarized 57 apple-breeding
tage over aneuploid or unbalanced gametes. projects in 25 countries (Ballard, 1998). Laurens
Researchers in Japan, Russia and the Ukraine (1999) surveyed apple-breeding programmes
internationally and summarized the primary 3.2.1.3 New York

objectives for scion cultivar improvement.
Cornell University’s apple-breeding pro-
Forty-two breeders from 29 countries
gramme at the New York State Agricultural
responded to the survey. The most common
Experiment Station in Geneva started in 1895
objective is to combine in new cultivars high
and has released 63 apple cultivars.
fruit quality with disease and pest resistance.
Scab (Venturia inaequalis (Cooke) Wint.) and ‘Cortland’, ‘Macoun’, ‘Empire’ and
powdery mildew (Podosphaera leucotricha (Ell. ‘Jonagold’ are among its best-known
& Ev.)) resistance is stressed. Programmes releases. The scab-resistant cultivar ‘Liberty’
develop cultivars adapted to an area’s extreme was released in 1978 and ‘Freedom’ was
climatic conditions. Genetic improvement of released in 1983. Brown and Terry (1997)
plant form involves selection of tree habits that reviewed the objectives of this programme in
allow high productivity and regular bearing. relation to traditional breeding, molecular
Hybridizations involve important commercial markers and biotechnology.
cultivars, old local cultivars and wild Malus
species. 3.2.1.4 Ohio
Mitch Lynd, a commercial grower, received
3.2.1 USA seedlings from germplasm collected in
Kazakhstan and planted them on his farm in
In 1983 there were 19 public apple-breeding 1999. This interest in disease-resistant, late-
and genetic programmes in the USA (Brooks blooming, hardy material led to the develop-
and Vest, 1985). Today there are only two ment of the Midwest Apple Improvement
full-time programmes – one each in Association, a grower group dedicated to
Washington State and New York. Five other the development of suitable cultivars for the
programmes in the USA divide their efforts Midwest. Additional information is at
between apples and other crops. http://www.hort.purdue.edu/newcrop/maia
/default/html
3.2.1.1 Arkansas
3.2.1.5 Purdue, Rutgers, Illinois (PRI)
Rom and Moore (1994) reviewed the pro- cooperative
gramme at the University of Arkansas,
where the development of cultivars to sup- The accomplishments of this cooperative pro-
port and expand the southern fruit industry gramme were reviewed by Crosby et al.
was an objective. ‘Stellar’ and ‘Arkcharm’ (1992). Recent scab-resistant releases include
were two of the cultivars released. ‘Goldrush’ and ‘Enterprise’ in 1993, ‘Pristine’
Hybridizations have stopped, but advanced in 1995 and ‘Scarlet O’Hara’ in 2000. Purdue
material continues to be evaluated. University is no longer making apple crosses,
but selections are still being tested (see
http://www.hort.purdue.edu/). Research at
3.2.1.2 Minnesota
the University of Illinois emphasizes the
Fruit breeding started at the University of improvement of apple by biotechnology.
Minnesota in 1878. Objectives are to develop
cold-hardy, high-quality, disease-resistant
3.2.1.6 New Jersey
cultivars. Noteworthy introductions include
‘Haralson’ in 1923, ‘Regent’ in 1963, ‘Sweet Rutgers University has named nine apple
Sixteen’ in 1979, ‘Honeycrisp™’ in 1991 and cultivars, with 11 additional cultivars
‘Zestar’ in 1999. The occurrence of several released as part of the PRI cooperative. Well-
episodes of low temperatures, from 30 to known releases include ‘Jerseymac’ in 1961,
38°C, in the winter of 1995/96 provided ‘Mollies Delicious’ in 1966, ‘Vista Bella’ in
data on cold-hardiness in the USDA core 1974, ‘Summer Treat’ in 1981 and ‘Suncrisp’
germplasm collection (Luby et al., 1999). in 1992. Fruit-ripening variants and their
effect on ethylene production is an area of ‘Belmac’ and ‘Primevere’ in 1996. A colum-

interest (Gussman et al., 1993). nar cultivar, ‘MacExcel’, was released for
the home market. In 1996, several partner-
ships with private industry were estab-
3.2.1.7 Washington
lished to test selected advanced apple
Washington State University started apple selections in commercial orchards in com-
breeding in 1994, with the goal of developing parison with commercially grown cultivars
cultivars adapted to the stressful hot, dry for their potential in the fresh market, long-
and sunny climate of central Washington term storage and juice and cider produc-
(Barritt, 1999). Seedlings are budded on to tion. A part of the programme is to select
M.9 and planted in an evaluation orchard 19 hardy disease- and insect-resistant cultivars
months later. Fruit evaluation began in 1999. for home gardeners. Several rootstocks,
with a range from vigorous to extremely
dwarf, are also under evaluation and might
3.2.1.8 Private breeders
be released for grower testing. Winter-
There are several private apple-breeding hardiness, pest and disease tolerance,
programmes in the USA. Two programmes dessert quality and long shelf-life remain
are part of commercial nurseries in breeding objectives. Additional information
California: Zaiger’s Genetics in Modesto and on the programme, selections and cultivars
Burchell Nursery in Oakdale. There are sev- is at http://www.pgris.com/
eral smaller programmes led by hobbyists or
commercial growers. 3.2.2.3 Nova Scotia, Kentville
This programme is known for the scab-
3.2.2 Canada resistant cultivars ‘Nova Easygro’ released in
1971, ‘Novamac’ in 1978 and ‘Novaspy’ in
3.2.2.1 British Columbia, Summerland 1986. Controlled hybridization has ended,
but the remaining seedlings and selections
Apple breeding started in 1924. Cultivars are being evaluated for potential release.
released for commercial use include Deslauriers et al. (1999) used descriptive sen-
‘Spartan’ in 1936, ‘Summerred’ in 1964, sory analysis (DSA) to define ten visual
‘Shamrock’ in 1986 and ‘Sunrise’ in 1991. In properties, nine flavour attributes and nine
1996, ‘Creston’ (‘Golden Delicious’ × an texture characteristics in apple and these
advanced selection), ‘Chinook’ (‘Splendour’ were used to evaluate selections for the fresh
× ‘Gala’) and ‘Silken’ (‘Honeygold’ × and processing markets.
‘Sunrise’) were named (Hampson et al.,
2000b). Two columnar or reduced branching
cultivars, ‘Scarlett Sentinel’ and ‘Golden 3.2.2.4 Manitoba, Morden
Sentinel’, were released in 1997 for the home The Dominion Experiment Station in
gardener. The Summerland programme Morden has as an objective the develop-
emphasizes determining consumer prefer- ment of new varieties of trees and shrubs
ences and sensory testing for quality evalua- and evaluation of material for adaptability
tion (Cliff et al., 1998; Hampson et al., 2000a). and value to the Canadian landscape trade.
Their website is http://www.em.agr.ca/ This programme released the ornamental
crab apples ‘Almey’, ‘Sundog’, ‘Garry’,
‘Selkirk’ and ‘Kelsey’, also known as the
3.2.2.2 Quebec
‘rosybloom’ crab apples. Scion cultivars
This programme originated in Ottawa in released include ‘September Ruby’, ‘Fall
the 1940s and was transferred to St-Jean- Red’ and ‘Red Sparkle’. Material developed
sur-Richelieu in 1971. Releases include at Morden can be grown in almost any loca-
‘Blair’ and the scab-resistant cultivars tion in Canada, due to the hardiness
‘Rouville’ and ‘Richelieu’ in 1990, and emphasis of the programme.
3.2.3 Australia combine durable resistance to scab, powdery

mildew and canker with fruit quality and a
3.2.3.1 Western Australia range of desirable horticultural characteris-
tics (Lateur et al., 1999, 2000).
Cripps Pink (‘Pink Lady®’) and Cripps Red
(‘Sundowner™’) were named in 1985 by the
Western Australian programme (Cripps et 3.2.5.2 Heverlee
al., 1993). Both cultivars are hybrids of
At the Fruitteeltcentrum at Katholieke
‘Golden Delicious’ × ‘Lady Williams’, a
Universiteit Leuven there are programmes in
long-storing cultivar believed to be a hybrid
classical breeding (started in 1989), transfor-
of ‘Granny Smith’ × ‘Rokewood’ (parentage
mation, haploid induction and molecular
unknown). This programme was broadened
understanding and characterization of self-
in 1985 and started producing 12,000
incompatibility. ‘Merlijn’, a hybrid of ‘King
seedlings per year until 1993. There are
Jonagold’ × ‘Liberty’, was the first release.
many advanced selections. In 1998 the goal
‘Merlijn’ has low susceptibility to scab and
was to maintain 50,000 seedlings.
mildew. More information on these diverse
Information on this national programme is
projects is at http://www.agr.kuleuven.ac.be/
at http://www.agric.wa.gov.au/ A focus
group, comprised of market representatives
from Asia, Europe and Australia, provides
3.2.6 Brazil
feedback on the market potential of
advanced selections.
Apple breeding at the Empressa de Paequisa
Agropecuaria e Extensao Rural de Santa
3.2.3.2 Queensland Catarina S.A. Experiment Station of Cacador
in Santa Catarina started in 1972 with seeds
Breeding at the Queensland Horticulture
imported from America. A low chilling
Institute in the Department of Primary
requirement for local adaptation and disease
Industries (DPI) at the Applethorpe Research
resistance are goals. ‘Princesa’ and ‘Primicia’
Station in Stanthope was started in 1964 to
(scab-resistant) were released in 1986, the
breed high-quality early red apples. An
low-chill, scab-resistant ‘Fred Hough’ was
emphasis on scab resistance was added in
named in 1994 and ‘Catarina’, a hybrid of
1986. ‘Applethorpe Earlidel’ was released in
‘Fuji’ × a scab-resistant selection, was named
1988 and ‘Applethorpe Summerdel’ in 1989.
in 1996. ‘Imperatiz’ and ‘Baronesa’ were
The website is http://www.dpi.qld.gov.au/
released in 1997. In 1997, two low-chill culti-
vars were released: ‘Condessa’, with more
scab and mildew resistance than ‘Princesa’,
3.2.4 Belarus
and the scab-resistant ‘Duquesa’. In 1999,
three cultivars adapted to regions with
Kozolovskaya et al. (2000) reviewed the 70-year
100–500 chilling units or less were released:
history in apple breeding, with 29 cultivars
‘Eva’, ‘Anabela’ and ‘Caricia’. These are also
released. Resistance breeding began in 1984
resistant to necrotic leaf blotch (Glomerella
and has yielded 136 progenies and over 30,000
spp.) and ‘Caricia’ is scab-resistant.
seedlings. Over 100 selections have been prop-
agated and several elites may be released.
3.2.7 Bulgaria
3.2.5 Belgium
Apple breeding started in the 1970s at the
Fruit Growing Institute in Plovdiv.
3.2.5.1 Gembloux
Objectives include early- and late-ripening
The programme at the Department of apples with disease resistance and a range of
Biological Control and Plant Genetic tree habits: standard, compact, columnar or
Resources started in 1988. Its goals are to weeping (Djouvinov, 1994).
3.2.8 China 3.2.8.5 Shandong Province

The Shandong Fruit Research Institute is at
Many of the provincial fruit-research insti-
Tai-an.
tutes in China have their own apple-
breeding programmes, resulting in over 180
new cultivars being released from 1950 to 3.2.8.6 Xinjiang Autonomous Region
1995. Of these, the majority resulted from
The Kueitun Fruit Research Institute released
controlled hybridizations, eight were from
‘XinShuai’ in 1985.
open pollination, two were induced muta-
tions and 11 were sports. ‘Ralls Janet’ is
popular in China and is in the pedigree of
3.2.9 Czech Republic
many releases.
The programme at the Research and
3.2.8.1 Hebei Province Breeding Institute of Pomology in
Holovouzy began in 1951. It has emphasized
The Hebei Changli Fruit Research Institute the use of polygenic resistance to scab from
released ‘Yanshanhong’, a hybrid of ‘Ralls old or lesser-known cultivars. Disease resis-
Janet’ × ‘Richared Delicious’, in 1989. tance and modification of plant form are
stressed. Six cultivars with scab tolerance
3.2.8.2 Henan Province have been named: ‘Angold’, ‘Julia’, ‘Klara’,
‘Nabella’, ‘Produkta’ and ‘Zuzana’ (Blazek
The Zhengzhou Fruit Research Institute of and Paprstein, 1994). Other releases impor-
the Chinese Academy of Agricultural Science tant in the Czech Republic include ‘Jarka’,
(CAAS) in Zhengzhou released two selec- ‘Julia’, ‘Resista’ and ‘Selena’.
tions in 1988: ‘Huaguan’ (‘Golden Delicious’
× ‘Fuji’) and ‘Huashuai’ (‘Fuji’ ×
‘Starkrimson’). Both have been noted for 3.2.10 England
their quality and storage life.
This programme emphasizes high quality,
long storage, the ability to compete with
3.2.8.3 Liaoning Province
imported fruit and resistance to mildew and
There are three programmes in this province, apple scab. Apples similar to ‘Cox’s Orange
with the Liaoning Fruit Research Institute Pippin’ were emphasized, but now diverse
having the largest apple-breeding pro- types are sought. Ten cultivars have been
gramme in China. One of their objectives is named since 1971. ‘Fiesta’, a ‘Cox’s Orange
to produce a new cultivar with all the advan- Pippin’ × ‘Idared’ hybrid released in 1984, is
tages of ‘Fuji’ but none of the disadvantages. well known. ‘Saturn’ (PRI 1235 × ‘Starkspur
‘Golden Delicious 463’, a russet-resistant Golden Delicious’) is a recent release. Six
induced sport, is their best-known introduc- columnar (reduced branching) cultivars have
tion. Shenyang Agricultural University been named: ‘Maypole’, ‘Telamon’, ‘Tuscan’
released ‘Hanfu’, a hybrid of ‘Dongguang’ × and ‘Trajan’ (Tobutt, 1985) and ‘Charlotte’
‘Fuji’. This cultivar is known for its hardiness and ‘Obelisk’ released in 1991. Molecular
and high eating quality. The Research markers are being developed for pre-
Institute of Pomology (CAAS) released selection at a juvenile stage (Evans, 1999).
‘Qiojin’ in 1975.
3.2.11 Finland
3.2.8.4 Shaanxi Province
The Shaanxi Fruit Research Institute at Breeding is conducted at the Agricultural
Yangling released ‘Quinguan’, a hybrid of Research Centre Institute of Horticulture in
‘Golden Delicious’ × ‘Jiguan’, in 1970. Pikkio.
3.2.12 France 3.2.13 Germany
3.2.12.1 Institut National de la Recherche 3.2.13.1 Ahrensburg

Agronomique, Angers
Apple breeding at the Federal Centre for
Objectives include resistance to pests and Breeding and Research on Cultivated Plants
diseases, reduction of orchard labour started in 1976, but was moved to
requirements and improved fruit quality. Dresden–Pillnitz in 1999 (see below). The
Several cultivars have been released for pro- goal was to develop high-quality cultivars
cessing and juice production, including with resistance to scab, mildew and Nectria
‘Jurella’, ‘Judaine’, ‘Judor’ and the scab- canker. Releases include ‘Ahrina’ (1989) and
resistant ‘Chanteline’. Other releases include the scab-resistant (Vf ) cultivars ‘Gerlinde’,
INRA Belchard® ‘Chantecler’ (‘Golden ‘Ahrista’ and ‘Ahra’. ‘Ahra’ also has low sus-
Delicious’ × ‘Reinette Clochard’), INRA ceptibility to canker.
Querina® ‘Florina’, a complex hybrid that is
resistant to scab and to rosy apple aphids, 3.2.13.2 Dresden–Pillnitz
released in 1985, and INRA ‘Baujade’, a
scab-resistant green apple with ‘Granny Apple breeding started in 1928 in
Smith’ in its pedigree, released in 1992. Muncheberg and was moved to
‘Initial’ is a new triploid, scab-resistant culti- Dresden–Pillnitz in 1971. The ‘Pi’ cultivar
var from ‘Gala’ × ‘Redfree’ that ripens about series (‘Pi’ for Pillnitz) were bred for high
1 week before ‘Gala’ (Laurens et al., 2000). quality, regular high yields and low suscepti-
INRA Perpetu® ‘Evereste’, an ornamental bilities to diseases. Ten ‘Pi’ cultivars have
crab apple, has also been released. Apple been named, including ‘Piros’, ‘Pilot’, ‘Pingo’
and ‘Pirella’. ‘Corail’, originally named
breeding at INRA also emphasizes the
‘Pinova’, a hybrid of ‘Clivia’ × ‘Golden
genetic modification of plant form. The aim
Delicious’, is of interest in Europe and in the
is to have a branching pattern that balances
USA. The ‘Re’ series is characterized by good
vegetative and reproductive growth, reduc-
fruit quality, high yield and different sources
ing pruning costs and reducing the tendency
of scab resistance. Fourteen ‘Re’ cultivars
towards biennial bearing (Lauri et al., 1997).
have been named: ‘Remo’, ‘Rewena’, ‘Rebella’
and ‘Reanda’ have resistance to scab, mildew
3.2.12.2 Commentry and fire blight (Fischer, 2000a,b).
Delbard International Nursery has a private

breeding programme. Cultivars released 3.2.14 Greece
include: ‘Delbarestivale® Delcorf ’ (‘Jongrimes’
× ‘Golden Delicious’), ‘Delbard® Jubilee’ The National Agricultural Research
(‘Golden Delicious’ × ‘Lundbytrop’), ‘Cybele® Foundation Pomology Institute in Naoussa is
Delrouval’ (‘Delcorf ’ × ‘Akane’), ‘Delearly’ incorporating both monogenic and polygenic
(‘Goldspur’ × ‘Stark’s Earliest’) and resistance. The local cultivar ‘Firiki’ has been
‘Harmonie® Delorina’ (‘Grifer’ × ‘Florina’). used in breeding. ‘Naoussa’ and ‘Makedoni’
Two of their cultivars are also being marketed were released from crosses made in 1973.
in the USA. ‘Delblush’ (cv. ‘Grifer’) is a
‘Golden Delicious’ × ‘Blushing Golden’ hybrid
called ‘Tentation’ in France. It received US 3.2.15 Hungary
Plant Patent (USPP) no. 10,276. ‘Regali®
Delkistar’ (‘Kidds’ Orange Red’ × ‘Bisbee Red Apple breeding started in 1972 at the
Delicious’) received USPP no. 11,213. University of Horticulture and Food
Industry in Budapest. The programme was 3.2.17.4 Ferraro

modified in 1984 and 1991 (Toth et al., 1994).
Private breeding by the nursery consortium
Quality, local adaptation and resistance to
Consorzio Italiano Vivaisti (CIV) has
multiple diseases and pests are priorities.
Four hybrids of ‘Jonathan’ × ‘Egri Piros’ resulted in the naming of ‘Rubens’ and
were released from 1985 to 1993. ‘Giotto’, both hybrids of ‘Gala’ × ‘Elstar’.
3.2.16 India 3.2.18 Japan
Sharma and Kumar (1994) reviewed fruit- Bessho et al. (1993) reviewed the status of
crop improvement in India and detailed 12 apple breeding and genetic analysis in
apple cultivars released. They suggested that Japan. The apple industry in Japan devel-
the limited success of fruit breeding in India oped following the importation of 75 culti-
might be due to the lack of continuity, spe- vars from America in 1871. ‘Ralls Janet’ and
cific gene sources and misdirected strategy. ‘Jonathan’ were two of the cultivars
imported. Apple breeding started at the
Aomori Apple Experiment Station in 1928.
3.2.17 Italy In 1939 a programme was started at the
Morioka Branch of the Fruit Tree Research
3.2.17.1 Bologna Station. There are seven prefectural research
stations and a national research station con-
The breeding programme at the University
ducting apple breeding.
of Bologna was initiated in 1976 and scab
Research in Japan focuses not only on
resistance was added in 1981. Objectives
cultivar development, but on all facets of
emphasize scab resistance, low susceptibility
genetic improvement. Resistance to
to mildew, spur or compact habit and high
Alternaria blotch has been identified and the
fruit quality (Sansavini and Ventura, 1994).
inheritance detailed. Resistance to apple
‘Prime Red’, a hybrid of ‘Prima’ ×
‘Summered’, was named in 1999. chlorotic leaf-spot virus (ACLSV) and apple
stem-pitting virus (ASPV) was determined
to be controlled by two recessive genes.
3.2.17.2 Forli Other studies have provided information on
The Instituto Sperimentale per la Frutticoltura the inheritance of tree habit, fruit colour,
programme at Forli started in 1980 to develop prevalence of burr knots and Valsa canker
improved dessert cultivars with resistance to resistance (Bessho et al., 1993).
scab and well adapted to the Po Valley.
3.2.18.1 Aomori Prefecture
3.2.17.3 Trento The Aomori Apple Experiment Station in
Apple breeding at the Trento section of the northern Japan has released many cultivars,
Experimental Institute of Fruit Culture started including the self-compatible cultivar
in 1974. Quality and pyramiding of resistance ‘Megumi’ in 1950 and ‘Mutsu’ in 1949,
are important objectives. Seven scab-resistant ‘Sekaiichi’ in 1974, ‘Tsugaru’ in 1975,
cultivars were released from 1997 to 1999: ‘Kitanosachi’ in 1979, ‘Hokuto’ and
‘Giongo’, ‘Red Earlilib’, ‘Golden Mira’, ‘Natsumidori’ in 1983 and ‘Mellow’ in 1990.
‘Brina’, ‘Nova’, ‘Summerfree’ and ‘Golden ‘Aori 9’, a hybrid of ‘Akane’ × ‘Orin’, which
Orange’. The last three cultivars are all is reported to be self-thinning, will be named
hybrids of PRI 1956-6 × ‘Ed Gould Golden’. (Kon et al., 2000).
3.2.18.2 Akita Prefecture 3.2.18.8 Nagano Prefecture

The Akita Fruit Tree Experiment Station The Nagano Fruit Tree Experiment Station
named ‘Senshu’ in 1983 and ‘Akita Gold’ established apple breeding in 1970. Releases
(‘Golden Delicious’ × ‘Fuji’) in 1990. include ‘Takane’ in 1982 and Nagano Apple
(NA) selections NA-10 (‘Fuji’ × ‘Tsugaru’),
NA-12 (‘Tsugaru’ × ‘Vista Bella’) and NA-15
3.2.18.3 Gunma Prefecture
(‘Golden Delicious’ × ‘Senshu’). Scab-
The Gunma Agricultural Research Centre resistance breeding was added in 1988.
released two hybrids of ‘Golden Delicious’ ‘Akane’ and ‘Alps-Otome’, a sweet crab
by an unknown pollen parent: ‘Akagi’ apple, are sources of scab resistance.
released in 1973 and ‘Yoko’ released in 1981.
They also named ‘Sinsekai’, a hybrid of ‘Fuji’
3.2.18.9 Private breeders
× ‘Akagi’, in 1988, and ‘Gunma Meigetsu’, a
hybrid of ‘Akagi’ × ‘Fuji’, in 1991. There are also about ten private breeders in
Japan who have released many cultivars,
including: ‘Akibea’, ‘Alps Otome’, ‘Kanki’
3.2.18.4 Fukushima Prefecture
(USPP no. 11,508), ‘Kinsei’, ‘Kogetsu’,
Breeding at the Fukushima Fruit Tree ‘Michinoku’, ‘Miki Life’ (USPP no. 11,511),
Experiment Station started in 1987, with the ‘Orin’ and ‘Scarlet’.
goal of developing cultivars suited to Japan’s
warmer areas.
3.2.19 Korea (South)
3.2.18.5 Hokkaido Prefecture
Breeding and biotechnology of apple culti-
The Hokkaido Prefectural Agricultural vars and rootstocks is overseen by the
Experiment Station released ‘Hacnine’, a National Horticultural Research Institute,
hybrid of ‘Fuji’ × ‘Tsugaru’, in 1984. Rural Development Administration. Apple
breeding is carried out at the Taegu Apple
Research Institute. Cultivars released
3.2.18.6 Iwate Prefecture
include: ‘Hwahong’, a hybrid of ‘Fuji’ ×
The Iwate Horticultural Experiment Station ‘Sekaiichi’, which ripens late in the season
released ‘Kio’, a hybrid of ‘Orin’ × and has a long storage life; ‘Hongro’ (‘Spur
‘Hatsuaki’, in 1992. Earliblaze’ × ‘Spur Golden Delicious’),
named in 1988; ‘Kamhong’ (‘Spur Earliblaze’
× ‘Spur Golden Delicious’) and ‘Chukwang’
3.2.18.7 Morioka
(‘Fuji’ × ‘Mollie’s Delight’), selected in 1992;
Apple breeding started in 1939 at the Apple and ‘Seokwang’, selected in 1996, an early-
Research Centre at the National Institute of season apple with an attractive colour and
Fruit Tree Science (NIFTS). Sixteen cultivars aromatic flavour.
have been named since 1952. Fruit quality,
storage ability and disease resistance are high
priorities. There have been three stages to the 3.2.20 Latvia
programme. ‘Fuji’ released in 1962, ‘Akane’ in
1970 and ‘Hatsuaki’ in 1976 were first named. The Dobele Horticultural Plant Breeding
The second phase resulted in ‘Kitakami’ in Experimental Station is studying local
1981, ‘Himekami’ in 1985 and ‘Iwakami’ in genetic resources and local cultivars that
1985. The third programme released ‘Sansa’ in may have been influenced by western and
1986 and ‘Kizashi’ in 1991. ‘Chinatsu’, a Russian cultivars. Twenty-eight cultivars
hybrid of ‘Akane’ × ‘Earliblaze’, and ‘Kitaro’, have been analysed for their breeding value.
a hybrid of ‘Fuji’ × ‘Hatsuaki’, were released The area is limited by low temperature peri-
in 1997. ‘Kotora’, a sib of ‘Kitaro’, was named ods of −30 to −35°C and by a short growing
in 1998 (Soejima et al., 2000). season of only 135–140 days (Ikase, 1999).
3.2.21 Lithuania produced annually. ‘Joy’ and Festival’ were

selected in the 1970s and commercialized in
Breeding at the Lithuanian Institute of several countries. Selections from ‘Gala’ ×
Horticulture in Kaunas also involves study ‘Splendour’ have been tested extensively in
of the inheritance of tree characteristics New Zealand and are being evaluated under
(height, diameter, branch number and non-distribution agreements in several coun-
internode length) and winter-hardiness tries. These include ‘Pacific Rose’ (Plate 3.3),
(Gelvonauskis, 1999). ‘Southern Snap’, ‘Sci Early’ and ‘Sci Red’
(Plate 3.4). Disease and insect resistance are
also being stressed (White and Bus, 1999).
3.2.22 Mexico Information on mapping, transformation,
disease and insect resistance and germplasm
The breeding programme at the Colegio de development is found at http://www.
Postgraduados, Fruticultura–IREGEP (Insti- hortresearch.cri.nz/
tute of Genetic Resources and Productivity)
has as an objective the development of low-
to medium-chill cultivars. 3.2.25 Norway
Breeding at the Ullensvang Research Centre

3.2.23 The Netherlands in Hermanswerk emphasizes the develop-
ment of improved early-ripening cultivars.
Breeding is at the Centre for Plant Breeding Four hybrids of ‘Katja’ × ‘Buckley Giant’ were
and Reproduction Research (CPRO-DLO) in introduced: ‘Nanna’, ‘Siv’, ‘Eir’ and ‘Idunn’.
Wageningen. From crosses made between
1948 and 1963, six selections were named,
with ‘Elstar’ being the most important. From 3.2.26 Poland
1964 to 1988 crosses, ‘Elan’ and ‘Elise’ CPRO®
were named. Crosses in the late 1970s 3.2.26.1 Skierniewice
emphasized scab resistance and resulted in Zurawicz and Zagaja (1999) reviewed breed-
the naming of the scab- and mildew-resistant ing at the Research Institute of Pomology
cultivar ‘Ecolette CPRO®’, a hybrid of ‘Elstar’ and Floriculture at Skierniewice. Three culti-
× ‘Prima’, in 1995. The next release, ‘Santana vars have been released: ‘Fantazja’
CPRO®’, a hybrid of ‘Elstar’ × ‘Priscilla’, is (‘McIntosh’ × ‘Linda’), ‘Alwa’ (‘Macoun’
resistant to scab and has good resistance to open-pollinated), and ‘Lodel’ (‘Lobo’ × ‘Red
European canker but is very susceptible to Delicious’). Two selections, ‘Redkroft’ and
mildew. A sweet apple from ‘Idared’ × ‘Ligol’ (‘Linda’ × ‘Golden Delicious’), are in
‘Elstar’ will be named ‘Bellida’. The web final testing.
address is http://www.plant.wag-ur.nl/
3.2.26.2 Warsaw
3.2.24 New Zealand This programme at the Warsaw Agricultural
University started in 1975. ‘Witos’, ‘Sawa’
McKenzie (1983) reviewed apple breeding and ‘Alka’ are cultivars that have been
from its ‘unofficial’ start in New Zealand released. ‘Sawa’, a hybrid of ‘Fantazja’ ×
over 100 years ago to the official start of a ‘Primula’, is considered to be a very good-
national programme at a government research quality, scab-resistant apple.
station in Havelock North in 1969. Research
scientists of the Department of Scientific and
Industrial Research produced three very 3.2.27 Romania
large families with 40,000 seedlings per fam-
ily. ‘Gala’, ‘Splendour’ and ‘Braeburn’ were Braniste (1997) reviewed apple breeding in
often parents of the 10,000–20,000 seedlings Romania, which started in 1948 in
Bucharest. Hybridizations are made in four Programmes also exist at the All-Russian
locations. Twenty-three cultivars have been Breeding and Technological Institute of
named, including ‘Frumos de Voinesti’, Horticulture and Nursery in Moscow and
‘Delios de Voinesti’, ‘Rosu de Cluj’ and the Research Institute of Horticulture of
‘Aromat de vara’. The scab- and mildew- Siberia in Barnaul.
resistant ‘Romus 1’, ‘Romus 2’ and ‘Romus
3’ were released in 1984 and ‘Pionier’,
‘Voinea’ and ‘Generos’ were named in 1985 3.2.29 South Africa (Republic of)
and 1986.
The Breeding and Evaluation Division of
Agricultural Research Council Infruitec/
3.2.28 Russia Nietvoorbij emphasizes the development of
high-quality, long-storing apple cultivars to
There are at least nine apple-breeding pro- give the South African industry a competi-
grammes in Russia, with many other pro- tive advantage. Development of locally
grammes being conducted as part of adapted cultivars is aided by studies of vari-
programmes within botanic gardens. While ation in winter chilling requirements and
most publications are in Russian, English prolonged dormancy symptoms. Breeding
abstracts of the breeding objectives and was started in 1955 and the apple cultivars
‘Drakenstein’ and ‘Gold Gift’ were released.
releases are available in some databases.
Scab-resistance breeding started in 1994.
Durable resistance to scab and powdery
‘African Carmine™’ is a 1999 release.
mildew, adaptation to adverse winter condi-
Information on the programme is available
tions, increased ascorbic acid content of the
at http://www.arc.agric.za/lnr/institutes/
fruit, reduced tree vigour and compact habit
niet/breeding/projects.html/
and greater use of local cultivars and wild
species are emphasized. It is difficult to
detail specific programmes, but some of the 3.2.30 Spain
locations include the following.
The Centro de Investigacion Aplicada y
3.2.28.1 The All-Russian Research Institute of Technologia in Villaviciosa has as an objec-
Horticultural Breeding in Orel tive the genetic improvement of cider apples.
Dr E.N. Sedov is the apple breeder. This

programme stresses high productivity, 3.2.31 Sweden
winter-hardiness, immunity to scab, high
ascorbic acid (vitamin C) content and breed- The Department of Horticultural Plant
ing for compact habit. Malus sylvestris, Breeding at the Swedish University of
Malus coronaria and ‘Kola’ crab apple have Agriculture in Balsgard released ‘Alice’ in
been used for improving vitamin C content. 1963, ‘Katja’, ‘Aroma’, ‘Sylvia’, ‘Kim’ and
The use of polyploids in breeding has been ‘Amorosa’ (‘Red Aroma’). Home-garden cul-
emphasized since 1970, especially crosses of tivars include the winter-hardy ‘Rodluvan’.
4x and 2x. Information on the programme is at
http://www.hvf.slu.se/ Objectives include
winter-hardiness, compact growth, good
3.2.28.2 The All-Russian Research Institute of
quality, resistance to diseases, high yield and
Horticulture (VNIIS) and All-Russian Research
long shelf- and storage-life.
Institute of Genetics and Breeding of Fruit
Plants in Michurinsk
The cultivar ‘Skala’ resulted from a cross of 3.2.32 Switzerland
I.V. Michurin’s cultivar × ‘Prima’. ‘Skala’ is
scab-resistant (Vf ) with large fruit and high The Swiss Federal Research Station in
vitamin C content. Wadenswil has released the cultivars
‘Maigold’, ‘Goro’, ‘Arlet’, ‘Iduna’ and to date. ‘Golden Delicious’ also has non-
‘Marina’ (Kellerhals and Meyer, 1994). Scab- russeting sports.
resistance breeding started in 1986 in cooper- To be most useful, mutations should be
ation with Horticulture Research periclinal mutations, a solid mutation that
International (HRI) in the UK and several includes all of LI and/or LII layers. If the
other institutions, and this has resulted in mutation includes the LII layer, where the
the naming of ‘Ariwa’, a scab- and mildew- gametes are formed, then the mutation will
resistant cultivar, bred in cooperation with also be useful in breeding. One example is
HRI. Information on the programme is at the columnar, or reduced branching, habit of
http://www.admin.ch/sar/faw/ ‘Wijcik McIntosh’, which is conferred by a
single dominant gene, Co (Lapins, 1974). This
sport is being used in crosses to study
3.2.33 Yugoslavia branching and to develop trees with modi-
fied plant form.
3.2.33.1 Novi Sad Sports that are initially mericlinal or sector-
ial can be stabilized. However, periclinal
The programme at the Faculty of
chimeras are sometimes unstable and may
Agriculture, Institute for Fruit Growing and revert back to the original cultivar, as exempli-
Viticulture started in 1985, with the breeding fied by the ‘MacSpur’ mutation of ‘McIntosh’
of columnar apples added in 1987 (Ognjanov (Embree et al., 1991). Pratt (1983) reviewed
et al., 1999). Objectives include preservation somatic selection and chimeras in fruit crops
of germplasm, breeding for multiple resis- and discussed the diploid–tetraploid sports in
tance using polygenic and monogenic apple and the development of a disbudding
sources and the production of cultivars technique to determine the inner phenotype of
suited for the home market, for use as polli- apple sports.
nators and as ornamentals. Mutation breeding of apples was once
popular, but declined in use because of the
3.2.33.2 Cacak high frequency of chimeras and undesir-
able or unstable forms (Lapins, 1983; Van
The Fruit and Viticulture Research Centre Harten, 1998). Mutation breeding empha-
at Cacak is part of the Agricultural sized compact types, non-russeting sports
Research Institute, Serbia. Apple breeding and sports with more fruit colour. Lacey
started in 1946. Since 1980 resistance breed- and Campbell (1987) reviewed the selec-
ing has been emphasized. Two hybrids of tion, stability and propagation of mutant
‘Starking Delicious’ × ‘Jonathan’ have been apples and the tests used to determine the
named: ‘Cacanska Pozna’ in 1971 and nature of the mutation. These methods of
‘Cadel’ in 1984. selecting and screening chimeras in muta-
tion breeding may also need to be used in
transgenic apple, where the possibility of
3.3 Natural and Induced Variation: chimeras of transformed and non-trans-
Mutations and Sports formed tissues must be considered (Ko et
al., 1998).
Apples are prone to limb or whole-tree White et al. (1994) studied red colour
mutations for enhanced fruit surface colour, changes following irradiation of ‘Royal Gala’
for spur-type growth habit and occasionally apple scions. They found a wide variation
for russet and ploidy differences (Pratt, among clones, but a small range of variation
1983). Cultivars differ in their tendency to within clones, consistent with the hypothesis
mutate, with ‘McIntosh’, ‘Delicious’, that the level of red colour expression is con-
‘Jonagold’, ‘Gala’ and ‘Fuji’ being very prone trolled by a multiple allelic series at the loci
to mutation. Yet ‘Empire’, derived from the governing red colour. Naturally occurring
frequently mutating ‘McIntosh’ and mutations of this multiple allelic series were
‘Delicious’, has only produced three sports proposed.
Concerns have been raised that redder 3.4.1 Heritability studies

sports appear to have a loss of flavour, in
contrast to the original parental clone. Heritability estimates provide breeders with
Fellman et al. (2000) reviewed the occurrence an indication of the potential for mass selec-
and biosynthesis of acetate esters, which are tion in improving a trait. Traits with high
the volatile compounds that impart the fruity heritability are more amenable to improve-
flavour and aroma associated with apples. ment, since the appearance (phenotype) is a
They found that higher-colouring strains of good indication of the breeding value (geno-
‘Delicious’ had lower aroma content. This type). Tancred et al. (1995) determined that
suggests that, when we select within a clone the heritability of ripening dates is high.
for higher pigmentation, quality may suffer. Durel et al. (1998) calculated narrow-sense
Molecular methods to differentiate among heritabilities for apple traits, using large
sports of a cultivar would be highly desirable unbalanced data sets. Heritability values
for the patenting, identification and protec- around 0.35–0.40 were obtained for fruit size,
tion of new sports. However, since the texture, flavour, juice content, attractiveness
genetic change is very small, current methods and russeting. Higher heritability values
have not proved useful. Pancaldi et al. (1999) were obtained for vigour, characterized by
used a randomly amplified polymorphic trunk circumference, and powdery mildew
DNA (RAPD) strategy called template mix- resistance (0.68). Currie et al. (2000) used 82
ing for the molecular analysis of mutant open-pollinated families to analyse fruit
clones, but this did not result in the additional shape and reported a combined-site heri-
bands (heteroduplexes) that are expected if tability of 0.79 for fruit aspect, best predicted
the primers are able to amplify the mutated by the fruit length/width ratio (R2 = 0.97).
DNA region. Tignon et al. (2000) used amplifi- Oraguzie et al. (2001) estimated narrow-sense
cation restriction-fragment length polymor- heritabilities for 19 traits in several sites,
phism (AFLP) to examine within-cultivar including fruit ribbing (0–0.13), fruit russet
variation, but found that it could not be used (0.05–0.58), fruit overcolour (0.34–0.40) and
for a secure identification of mutant cultivars. fruit weight (0.27–0.90).
3.4 Genetics and Inheritance of Traits 3.5 Disease Resistance: Present Status
and Future Directions
Alston et al. (2000) updated the Malus gene
list to include 145 genes, with 29 involved in 3.5.1 Apple scab (V. inaequalis (Cooke)
pest or disease resistance. Similar lists and Wint.)
discussions are found in Brown (1992) and
Janick et al. (1996). Gene discovery is increas- The discovery, characterization and use of
ing as sequences from other species are being genes for resistance to apple scab demon-
used to investigate Malus. strates the importance of diversifying
Genetic research on plant architecture, sources of resistance. Williams and Kuc
spurriness, bearing habit and regularity of (1969) summarized both qualitative and
production is also of high priority. Tobutt quantitative sources of resistance to apple
(1994) crossed cultivars with apetalous flow- scab. Although six genes for scab resistance
ers – ‘Wellington Bloomless’ and ‘Spencer were available, breeders concentrated pri-
Seedless’ – with four columnar selections marily on Vf . The Vm gene from Malus micro-
and determined that the gene ape for malus and Malus atrosanguinea 804 was
apetalous was recessive. Apetalous cultivars susceptible to race 5, genes from ‘Russian
tend to set fruit parthenocarpically. seedling R12740-7A’ (Vr and Vx) were
Apetalous parthenocarpic selections would known to have differential resistance and the
have a reduced need for pollination and, resistance gene Vbj from Malus baccata jackii
with less seed production, might be less was not used by many programmes. The
prone to biennial bearing. danger of concentrating on one source of
qualitative resistance became evident in 1993 vars. Information on DARE is available at

with the discovery of race 6. Race 6 induced h t t p : / / w w w. i n r a . f r. / A n g e r s / D A R E /
scab on nearly all Vf selections, but did not Polygenic resistance in ‘Antonovka’,
infect Malus floribunda 821, one of its selected ‘Discovery’, ‘James Grieve’, ‘Dulmener
F2 progeny (F2 26829-2), ‘Granny Smith’, M. Rosenapfel’ and the old Italian cultivars
baccata jackii (Vbj) and ‘Russian seedling ‘Durello di Forli’ and ‘Renetta Grigoa di
R12740-7A’ (Vr, Vx) (Parisi and Lespinasse, Torriana’ will be studied.
1999). The discovery of race 7 and the inter-
action between race 6 and race 7 with the
original source of resistance, M. floribunda 3.5.2 Powdery mildew (P. leucotricha (Ell. &
821, suggested the existence of a second Ev.) Salm.)
dominant gene, independent of Vf (Benaouf
and Parisi, 2000). This gene was named Vfh Many commercial cultivars being used in
because it induced a hypersensitive breeding, such as ‘Ginger Gold’,
response. This finding supports theories that ‘Honeycrisp’ and ‘Goldrush’, are highly
Vf resistance was more complex than it being susceptible to powdery mildew, making
dependent only on a single gene. Results parental and progeny selection for resis-
from race 7 identified another dominant tance important. Breeding for resistance to
gene, Vg, responsible for the resistance of powdery mildew has been difficult due to a
‘Golden Delicious’ to strain 7. poor understanding of the races and their
Breeders need to pyramid sources of geographical locations and often a poor cor-
resistance and to use molecular markers in relation between seedling (i.e. juvenile
pyramiding to ensure that more than one plant) reaction to infection and that of
gene for resistance is present. We need to mature plants.
have a better understanding of differential There are both qualitative and quantitative
reactions (Gessler and Blaise, 1994) and of sources of resistance in Malus. Knight and
the resistance in ‘Antonovka’ (Va and poly- Alston (1968) first proposed single gene con-
genic) and in ‘Jonsib’. We need to character- trol of resistance, but later two different genes
ize and use polygenic or partial resistance plus modifiers were proposed: Pl1 from Malus
and/or the natural low susceptibility of old × robusta and Pl2 from Malus × zumi Rehd. In
cultivars, new cultivars and advanced selec- 1977, Dayton identified an open-pollinated
tions. The use of polygenic resistance will seedling from ‘Starking Delicious’ as highly
require modifications of existing protocols resistant. The seedling’s pollen parent was
and examination of factors such as the influ- believed to be a Malus species. This seedling
ence of scab inoculum concentration in was designated mildew-immune seedling
screening for quantitative resistance (Lateur (MIS). Alston et al. (2000) proposed the desig-
et al., 2000). nation Pl-m for this resistance. Alternatively,
Durable Apple Resistance in Europe Korban and Riemer (1990) examined segrega-
(DARE) is a project funded by the European tion among 14 progenies for mildew reaction
Union to detect and characterize durable and indicated that mildew resistance was
sources of resistance in local European culti- polygenically controlled in this material with
vars. Nine institutions participate in the additive gene effects.
project. Lespinasse et al. (2000) summarized
the work in progress. DARE focuses on
durable resistance to scab and powdery 3.5.3 Fire blight (Erwinia amylovora (Burrill)
mildew. Goals include: (i) characterization Winslow)
of resistance of cultivars; (ii) assessment of
the pathogenicity and variability of the two Many popular scion cultivars and rootstocks
fungi; (iii) genetic dissection of partial resis- are highly susceptible to this bacterial
tance; (iv) development of new breeding pathogen. Therefore, the use of genetic resis-
strategies; and (v) market study and con- tance by traditional breeding and by biotech-
sumer preference for new resistant culti- nology is a goal of many programmes. The
scion cultivar ‘Liberty’ is a good source of 3.6 Other Traits

resistance and may also be used as a control
cultivar to assess resistance in transgenic lines. 3.6.1 Quality
Gardner et al. (1980a) evaluated Malus
species and clones for resistance and found The improvement of fruit quality is one of
that the small-fruited Asian species, espe- the most important breeding objectives, but
cially Malus sieboldii, were outstanding genetic manipulation is hampered by the
sources of resistance. There were very few complexity of the topic, the influence of the
domestic cultivars with high resistance and environment and the lack of measurements
intraspecific variation was noted. They cau- that are quantitative rather than subjective.
tioned that screening and inoculation based The definition of quality will vary depend-
on young seedlings might not reflect the sus- ing on the programme and market require-
ceptibility of older shoots. The inheritance of ments, but flavour, texture (crispness and
resistance was studied. There was 90% mor- firmness) and juiciness are three components
tality in crosses of susceptible × susceptible. of interest to most breeders. Genetic
Resistant × susceptible yielded few resistant improvement of postharvest quality is also
offspring and resistant × resistant still pro- important.
duced some highly susceptible seedlings, Apple flavour, including taste, aroma, off-
suggesting quantitative control of resistance. flavours, primary and secondary volatiles
In crosses of the highly resistant M. × robusta and correlations between instrumental and
No. 5 × Malus × sublobota PI286613, resistance sensory analysis, was reviewed by Yahia
was suggested to be controlled by a few (1994). He suggested focusing on odour-
genes (Gardner et al., 1980b). Polygenic con- active volatiles as a key to future molecular
trol of resistance was also suggested in a manipulation.
study of eight interspecific and intraspecific
Many programmes are using sensory
crosses. Malus prunifolia var. xanthocarpa and
testing as part of their evaluations.
M. prunifolia var. microcarpa were good
Hampson et al. (2000a) detailed the use of
sources of resistance (Korban et al., 1988).
sensory evaluation as a selection tool in
Evaluation of greenhouse-grown seed-
apple breeding. They determined that a
lings inoculated with a mixture of strains of
minimum panel size of 11 was necessary to
the bacteria was found to be a better predic-
obtain statistical discrimination of one point
tor of field susceptibility than evaluation
on a zero-to-nine scale. Crispness was found
with single strains (Norelli et al., 1986). With
to account for 90% of the variation in texture
the discovery of differential susceptibility,
liking. Perceived sweetness and sourness
Norelli et al. (1986, 1987) cautioned that
progenies must be inoculated with strains were better predictors of liking than fruit
that are representative of the complete Brix and titratable acid. Deslauriers et al.
range of differential virulence when breed- (1999) used DSA to define and quantify the
ing for resistance. sensory properties of apple and used statis-
Chevreau et al. (1998) examined four tical programmes to select genotypes most
somaclonal variants of ‘Greensleeves’ for similar to a target cultivar, such as a popular
resistance and found that in vivo tests were processing type, but with enhanced produc-
unreliable due to high variability, that green- tivity. The use of sensory assessment also
house inoculation was more reliable but proved valuable in detecting quantitative
overrated resistance and that field inocula- trait loci (QTL) representing different attrib-
tion was the most severe test. Transformation utes of fruit texture (King et al., 2000).
of apple with lytic peptides to confer resis- Significant QTL were detected on seven
tance to fire blight has been successful linkage groups. Magness–Taylor penetrome-
(Norelli et al., 1999). Transgenic lines of ter readings, stiffness by acoustic resonance
‘Royal Gala’ and M.7 rootstock with the gene and sensory descriptors assessed by a
encoding attacin E resulted in a significant trained panel were the instrumental mea-
increase in resistance to fire blight. surements used.
3.6.2 Nutrition 3.7.1 Isozymes
Increasing the ascorbic acid or vitamin C Isozymes were the first markers to be used in
concentration in apples has long been a goal apple breeding. They have been used for
in apple breeding. While most cultivars identification of cultivars, for estimating
contain from 5 to 10 mg ascorbic acid g−1, genetic diversity in germplasm collections
cultivars with much higher concentrations and to confirm or refute suspected parent-
(20–50 mg g−1) have been documented. age. Chyi and Weeden (1984) used isozymes
While vitamin C concentration is important, to determine that the female parent supplied
the role of other antioxidants is also of the unreduced diploid gamete in several
interest and is likely to be a new area for triploid cultivars. Weeden and Lamb (1987)
genetic improvement. Eberhardt et al. (2000) studied isozyme polymorphism in nine
demonstrated that 100 g of fresh apples enzyme systems and identified four linkage
have an antioxidant activity equivalent to groups. Close linkage of glutamate oxaloac-
1500 mg of vitamin C. Apple phytochemi- etate transaminase (Got-1) and the S incom-
cals, phenolic acids and flavonoids signifi- patibility locus was reported (Manganaris
cantly enhance the antioxidant properties and Alston, 1987). Manganaris and Alston
and are amenable to genetic improvement. (1988) also found that the acid phosphatase
gene ACP-1 was linked with the endopepti-
dase gene ENP-1 and the pale green lethal
3.6.3 Reduction of flesh browning gene, l. In 1994, Manganaris et al. reported
that the isozyme locus Pgm-1 was tightly
Genetic reduction of apple flesh browning is linked to Vf scab resistance. Advances con-
also a breeding objective. For the fresh-cut tinue to be made. Alston et al. (2000)
slice market, cultivars will need to have low reviewed research on 69 isozymes in apple.
levels of polyphenol oxidase and sufficient
ascorbic acid and to maintain their crispness
and firmness. Transgenic approaches to 3.7.2 Scab resistance
reduced browning are also being tested.
The amount of research focused on the iden-
tification and development of markers for
3.6.4 Allergenicity the Vf gene is extensive. Only the most recent
studies will be discussed. Patocchi et al.
Research on the allergenicity of apples has (1999) used markers to fine-map the Vf
expanded. Puhringer et al. (2000) found that region. Xu and Korban (2000) used AFLP
the promoter for the major allergen Mal d1 markers for saturation mapping of Vf and
is stress- and pathogen-induced. Breeders then sequence-characterized amplified
need to ensure that new cultivars are not regions (SCARs) were developed from these
higher in allergenicity and need to explore AFLP markers (Xu et al., 2001). These are
prospects of breeding apples with lower prerequisites for map-based cloning.
allergenicity. Markers have also been developed for the
Vm region from M. atrosanguinea 804 (Cheng
et al., 1998). Research on markers for the
3.7 Markers and Genetics other genes for scab resistance is progressing
in several laboratories.
The ability to use molecular markers for pre-
selection at the seedling stage, prior to field
planting, is of great importance. The ability 3.7.3 Powdery mildew
to prescreen reduces land and greenhouse
requirements, saves money and results in a Molecular markers for the resistance genes
higher percentage of desirable seedlings Pl1 from M. × robusta (Markussen et al., 1995),
being planted in the field. Pl2 from M. × zumi Rehd. (Dunemann et al.,
1999) and Plw from ‘White Angel’ (Batlle and lines appeared to have the incompatibility
Alston, 1996) are being used in marker- mechanism switched off, as evidenced by
assisted selection. pollen-tube growth.
3.7.4 Insect resistance 3.7.6 Other traits
Markers have been developed for resistance Markers have been identified for many traits,
to the rosy leaf-curling aphid (Dysaphis including fruit skin colour (Cheng et al.,
devecta Wlk.) (Roche et al., 1997a,b) and to 1996), columnar habit (Hemmat et al., 1997)
woolly apple aphid (Eriosoma lanigerum and fruit acidity (malic acid concentration)
Hausmn.), an important pest in breeding (Conner et al., 1997). Conner et al. (1998) used
rootstocks (Bus et al., 2000). RAPDs to estimate the position and effect of
QTL affecting juvenile tree growth and
development and found that a large number
3.7.5 Incompatibility alleles of traits had significant variation associated
with the map position of the dominant
The self-incompatibility system in apple is columnar gene, Co.
well known, but until recently the number
of S alleles involved and the extent of cross-
incompatibility was not known. The cloning 3.7.7 Microsatellites/simple sequence
and molecular analysis of two self-incom- repeats (SSRs)
patibility alleles from apple (Broothaerts et
al., 1995) was followed by the development SSRs, short tandem repeats of one to six base
of a molecular method for S-allele identifica- pairs, have been used in cultivar identifica-
tion in apple based on allele-specific PCR tion and genetic analysis and to reveal iden-
(Janssens et al., 1995). The alleles S2, S3, S5, tities, genetic diversity and relationships in a
S7 and S9 were identified and used to geno- core subset collection. Guilford et al. (1997)
type several cultivars, including ‘Idared’ used SSRs in a survey of 21 cultivars. The
(S3S7), ‘Fiesta’ (S3S5), ‘Jonathan’ (S7S9), majority of SSRs were highly polymorphic
‘Elstar’ (S3S5), ‘Gala’ (S2S5) and ‘Golden and diploid and showed simple Mendelian
Delicious’ (S2S3). Sakurai et al. (1997, 2000) inheritance, although about 25% of markers
used this method to genotype Japanese and generated complex banding patterns. Three
American apple cultivars and advanced microsatellite markers were sufficient to dif-
selections. Six additional S alleles were ferentiate between all 21 cultivars.
sequenced: S4, S24, S26, S27, Sd and Sf Gianfranceschi et al. (1998) developed 16 SSR
(Sassa et al., 1996; Katoh et al., 1997; markers that amplified all alleles from 19
Verdoodt et al., 1998). S genotyping has cultivars, breeding selections and M. flori-
raised questions of paternity in that several bunda 821. Two selected SSRs were able to
cultivars have S alleles different from those distinguish all cultivars except ‘Starking’ and
predicted by their parentage. ‘Red Delicious’. Hokanson et al. (1998)
S-allele genotyping has also been used to screened accessions from a core subset of the
assess homozygosity in shoots obtained germplasm repository with eight SSRs. The
through haploid induction by screening in primer pairs differentiated all but seven
vitro shoots for single S alleles as opposed to pairs of accessions.
S alleles of a parent whose pollen was irradi-
ated and used to stimulate parthenogenic
development (Verdoodt et al., 1998). Van 3.7.8 Molecular maps
Nerum et al. (2000) transformed ‘Elstar’ with
an S allele in the sense or antisense direc- The first linkage map of apple was based
tion and analysed lines for self-fertility. on a progeny of 56 seedlings from a cross of
Fluorescent microscopy confirmed that some ‘Rome Beauty’ × ‘White Angel’ that com-
bined isozyme, RAPD and restriction- is a prerequisite for the construction of a

fragment length polymorphism (RFLP) physical map of apple and for map-based
markers (Hemmat et al., 1994). Conner et al. cloning of Vf or other apple genes.
(1997) developed maps for ‘Wijcik
McIntosh’ and for two advanced scab-resis-
tant selections from the Cornell breeding 3.8 Biotechnology
programme. Maliepaard et al. (1999)
reviewed the maps for ‘Prima’ × ‘Fiesta’, Name recognition in marketing apples is
the first progeny used for mapping apple in important. Thus, using biotechnology to
Europe. One hundred and fifty-five F1 change a key characteristic of a popular com-
seedlings were genotyped with 208 mark- mercial cultivar and yet maintain varietal
ers, which included RFLPs, RAPDS, isoen- identity is very desirable. This objective can-
zymes and microsatellites. The European not be achieved in traditional breeding
DARE programme has as one of its goals because of the need to use a modified back-
the identification of molecular markers cross procedure due to self-incompatibility
linked to genes for resistance. Five molecu- and inbreeding depression.
lar maps, mainly involving SSRs and
AFLPs, have been constructed. Research is
conducted in France, Germany, Italy, 3.8.1 Somaclonal variation
Greece, The Netherlands and Switzerland.
Mapping populations also include ‘Fiesta’ There has been an ongoing debate about the
× ‘Discovery’; ‘Fiesta’ is susceptible and effect of tissue culture on apple and the
‘Discovery’ has a high level of resistance to extent of somaclonal variation that might
both scab and mildew. Researchers at INRA exist. This has important implications for
in France are using the cross ‘Discovery’ × the genetic transformation and regeneration
TN10-8. Researchers at the University of of commercial cultivars. Somaclonal varia-
Bologna in Italy and in Greece are mapping tion for resistance to the fire blight
‘Durello di Forli’ × ‘Fiesta’, while those in pathogen E. amylovora and for alterations to
Ahrensberg, Germany, are mapping ‘Prima’ rooting ability and shoot proliferation in
× ‘Discovery’. A common set of AFLPs will vitro was examined by Donovan et al.
be tested in mapping populations that have (1994a,b). Zimmerman (1997) reported that
‘Discovery’ as a parent. micropropagated trees of ‘Redspur
Delicious’ exhibited tree-to-tree variation
and that most replicates did not maintain
3.7.9 Comparative mapping the spur habit. However, micropropagating
spur-type trees from previously microprop-
Mapping of resistance-gene analogues agated trees that did retain the spur habit
(RGAs) from other species is ongoing. was successful in having spur habit main-
Comparative mapping with other members tained. When tissue culture-derived ‘Gala’
of the Rosaceae, especially with Pyrus and and ‘Royal Gala’ clones that were obtained
Prunus, is likely. Several microsatellite via axillary and adventitious bud formation
repeats in peach (Prunus persica (L.) were compared with conventionally grafted
Batsch.) were also amplified in apple trees by McMeans et al. (1998), very little
(Cipriani et al., 1999). somaclonal variation was observed in mor-
phological or reproductive traits (Plate 3.5).
Yet Zimmerman and Steffens (1995)
3.7.10 Bacterial artificial chromosome (BAC) reported that tissue-cultured ‘Gala’ trees
library of apple often developed burr knots 6–7 years after
being transferred to the field. They sug-
Vinatzer et al. (1998) reported the construc- gested that tissue cultures should be re-
tion of a BAC library using ‘Florina’, a scab- established annually to prevent this
resistant cultivar (Vf gene). The BAC library problem.
3.8.2 Regeneration and transformation to the herbicide Glean™ in transgenic plants.

James et al. (1996) documented the stable
The literature in this area is extensive. The expression and Mendelian segregation of the
reader is referred to several reviews as a marker transgenes nopaline synthase (nos)
starting-point, but advances are continually and the cotransferred gene neomycin phos-
being made. Protoplast fusion (symmetric photransferase (nptII) in the flesh of apple
and asymmetric) has been tested in apple fruits 7 years after the initial transformation.
and some tentative somatic hybrids have In 1996, ‘Gala’, ‘Golden Delicious’ and
been identified (Huancaruna Perales et al., ‘Elstar’ were transformed (Puite and Schaart,
2000). Singh and Sansavini (1998) reviewed 1996), followed by ‘Delicious’ and ‘Pink
transformation across fruit crops, while Lady®’ (Sriskandarajah and Goodwin, 1998)
Hammerschlag (2000) reviewed transforma- and ‘Delicious’, ‘Greensleeves’ and ‘Royal
tion of Malus. De Bondt et al. (1994, 1996) Gala’ (Maximova et al., 1998).
reviewed factors influencing gene-transfer Bolar et al. (1999) developed an efficient
efficiency during early transformation steps transformation system for ‘Marshall
and factors affecting regeneration of trans- McIntosh’. Expression of endochitinase from
formants. Maximova et al. (1998) investi- Trichoderma harzianum in apple increased
gated transformation using green fluorescent resistance to scab and reduced vigour in
protein and found that high transient expres- transgenic ‘Marshall McIntosh’ (Bolar et al.,
sion and low stable transformation sug- 2000). There was a significant negative corre-
gested that factors other than (T)-DNA lation between the level of endochitinase
transfer were rate-limiting. production and both the amount of disease
and plant growth.
Yao et al. (1999a) grew transgenic ‘Royal
3.8.3 Transgenes introduced Gala’ apple trees under controlled green-
house conditions and 20% of the trees flow-
The first report of transformation of apple ered in the second year, but, when scion
occurred in 1989 (James et al., 1989). wood from the top of these clones was
Trifonova et al. (1994) transformed ‘Granny grafted on to M.9, 85% produced flowers and
Smith’ with nptII and ipt genes, encoding for fruit the next year. Inheritance of three trans-
one of the first enzymes in the cytokinin genes, uid A, neomycin phototransferase II
biosynthetic pathway (Fig. 3.1). In 1995, Yao and acetolactate, fit a 1 : 1 ratio in most lines,
et al. introduced the acetolactate synthase but in one progeny line the T-DNA integra-
gene into ‘Royal Gala’ to increase resistance tion pattern was complex.
Fig. 3.1. Machine used to transform apple tissue by inserting genes.

Two of four transgenic lines possessing clones rooted (83–100%) on hormone-free

the kanamycin resistance gene and antisense rooting medium versus 1% for the controls.
polyphenol oxidase (PPO) DNA showed Root length and root morphology did not
repressed PPO activity and a lower brown- differ between transgenic clones and the
ing potential than control shoots (Murata et untransformed controls.
al., 2000). Broothaerts et al. (2000b) developed
a spectrophotometric assay for the analysis
of PPO in apple and tobacco leaves to 3.8.5 Transgene silencing
increase efficiency in screening large num-
bers of transgenic plants (Fig. 3.2). Ko et al. (1998) found that there were alter-
Transformation of ‘Jonagold’ with antimi- ations in nptII and gus expression following
crobial peptide genes (A1-AMP) resulted in micropropagation of transgenic M.7 apple
28 independent transgenic lines, which are rootstock lines. The gus gene was present in
being tested for resistance to apple scab non-staining lines. Gus gene silencing was
using artificial inoculation assays due to methylation in some cases, but in
(Broothaerts et al., 2000a). At the Apple others the mixed staining might be due to a
Research Centre in Morioka, Japan, ‘Orin’ mixture of transformed and non-
and the Japanese rootstock ‘JM 7’ have been transformed cells.
transformed with genes encoding the sor-
bitol-metabolizing enzyme sorbitol-6-
phosphate dehydrogenase isolated from 3.8.6 Challenges
apple, chitinase isolated from rice, glucanase
from soybean and sacrotoxin from the flesh- At present, traits that are complex (e.g. yield
fly (Soejima et al., 2000). and flavour) are not likely candidates for
improvement by biotechnology. Additionally,
there is a need for genes from Malus to be
3.8.4 Rootstocks transformed cloned, since public concern about transgene
technology does differentiate between
Apple rootstocks are also a focus in biotech- native and non-native genes. There is also a
nology, with M.26 rootstock (Lambert and need for specific promoters, wound-
Tepfer, 1992; Maheswaran et al., 1992; inducible or fruit- or leaf-specific, so that
Holefors et al., 1998) and M.7 rootstock transgene expression may be targeted only to the
formed (Norelli et al., 1999). M.26 was also parts of the plant necessary for the desired
transformed with rolA and rolB (Zhu and effect (Gittins et al., 2000). Transgenic testing
Welander, 2000). Zhu et al. (2001) trans- must ensure that there are no non-target
formed M.9 rootstock with rolB and found effects and that transgenic lines are stable
that in in vitro rooting tests all transgenic and non-chimeric.
Fig. 3.2. Gel comparing banding patterns of seedlings segregation for disease resistance.
3.9 Future Prospects logue of genes for DAD1 (defender against

cell death 1). Southern hybridization indi-
3.9.1 Malus genes being investigated cates that apple contains at least two DAD1
homologues. Transcript levels vary between
As researchers use sequence-information tissues and are induced by flower pollina-
comparisons with Arabidopsis mutants and tion and senescence of leaves, petals and
other well-characterized plants, genes from fruits. MdDAD1 mRNA was distributed
Malus are being identified and cloned primarily in the vascular bundles. Other
(http://www.ncbi.nlm.nih.gov/). Research research identified apple mRNAs related to
on MADS in apple is an example of how bacterial lignostilbene dioxygenase and
initial findings may evolve. In 1997, a plant small-auxin-up-RNA (SAUR) genes,
MADS-box cDNA clone of the ‘Fuji’ apple which were preferentially expressed in
was cloned and characterized (Sung and flowers (Watillon et al., 1998). Kotoda et al.
An, 1997). Yao et al. (1999b) found that (2000) examined expression patterns of
MADS-box genes were expressed in differ- homologues of floral meristem identity
ent parts of the fruit. Parthenocarpic apple genes LFY and AP1 during flower develop-
fruit production was then found to be con- ment in apple. These studies should
ferred by transposon insertion mutations in advance our understanding of floral initia-
a MADS-box transcription factor (Yao et al., tion and development.
2001).
3.9.4 Summary
3.9.2 1-Aminocyclopropane-1-carboxylate
(ACC) synthase and ACC oxidase Advances in molecular markers, gene char-
acterization and sequencing and the trans-
In 1991, Dong et al. cloned a cDNA encoding formation and expression of transgenes
ACC synthase, the main gene responsible for offer great potential to aid the efficiency and
ethylene production during ripening. In effectiveness of genetic improvement in
1998, genomic clones associated with ACC apple. These advances will allow us to
oxidase and polygalacturonase mRNAs in manipulate genes affecting quality, disease
ripe apples were isolated and expression was resistance and plant architecture. Cloning
monitored in three cultivars (Dong et al., genes from Malus will occur with greater
1998). The activity and tissue specificity of frequency and perhaps aid our understand-
the promoters were analysed in transgenic ing of the role of transposons in the activa-
tomato (Atkinson et al., 1998). Harada et al. tion of mutations for colour and plant habit.
(2000) hypothesized that a low level of ethyl- Transformation using native genes may
ene production might be caused by a enhance our knowledge of gene silencing
mutated allele of the ACC synthase gene and perhaps gene activation. Adding genes
(Md-ACS1). for resistance to apple cultivars with native
resistance to disease is likely to impart
broader-scale resistance with less likelihood
3.9.3 Additional genes involved in flowering of resistance breakdown. Undoubtedly, new
information and new techniques will con-
Dong et al. (1998) constructed a cDNA tinue to lead to improvements in fruit qual-
library from developing fruits of apple, 2 ity, storage life, nutrition, resistance to
days after pollination. Differential screening insects and diseases and self-fertility, result-
for pollination-induced genes resulted in ing in new apple cultivars and improve-
the isolation of MdDAD1, an apple homo- ments in existing cultivars.
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Apples - Chap 04 21/3/03 2:55 pm Page 61
4 Characteristics of Important Commercial

Apple Cultivars
Cheryl R. Hampson1 and Henk Kemp2

1Pacific Agri-Food Research Center, Agriculture and Agri-Food Canada, Summerland,
British Columbia, Canada; 2Applied Plant Research, Fruit Section, Radwijk,
The Netherlands
4.1 Introduction 61
4.2 Cultivar Descriptions 62
4.2.1 ‘Delicious’ 62
4.2.2 ‘Golden Delicious’ 64
4.2.3 ‘Fuji’ 66
4.2.4 ‘Granny Smith’ 68
4.2.5 ‘Gala’ 69
4.2.6 ‘Jonathan’ 71
4.2.7 ‘Jonagold’ 72
4.2.8 ‘McIntosh’ 74
4.2.9 ‘Rome Beauty’ 76
4.2.10 ‘Braeburn’ 77
4.2.11 ‘Elstar’ 78
4.2.12 ‘Cox’s Orange Pippin’ 80
4.3 Outlook 81
4.1 Introduction (Janick et al., 1996). The high cost of mod-

ern production requires a cultivar to have
The cultivated apple is believed to have orig- prolific, consistent yields of uniform, com-
inated in central Asia. Its chief ancestor is mercial-quality fruit, be amenable to han-
probably Malus sieversii, from the Heavenly dling, storage and shipping and generate
Mountains (Tien Shan) on the border of high consumer demand. Resistance to dis-
western China, the former USSR and central eases, pests and storage disorders is also
Asia (Janick et al., 1996). Apples have been important. In the past, most small farms
cultivated and vegetatively propagated for produced their own apples for fresh or
over two millennia. preserved use and local markets.
Over 10,000 named cultivars exist and Improvements in storage technology elimi-
breeders worldwide create more new selec- nated the need for a succession of short-
tions annually, but only a few dozen types storing apples from early summer to late
are widely produced in commerce today winter (French, 1970) and displaced some

62 C.R. Hampson and H. Kemp
old, long-keeping apples with less con- 4.2.1 ‘Delicious’

sumer demand, e.g. ‘Winesap’. Improved
storage and year-round commercial avail- ‘Delicious’ arose unwanted in the field of
ability also shifted consumption patterns Jesse Hiatt of Peru, Iowa, USA, in about 1872
from preserved forms (cider, vinegar, dried (Maas, 1970a; Fear and Domoto, 1998). Mr
apples, apple butter, apple sauce) to fresh Hiatt cut the tree down twice before allow-
apples. Societal changes, including the tem- ing it to persist, because it was not growing
perance movement, reduced the demand in the row. The parentage is unknown; a
for cider apples. Apples grown by farm nearby ‘Yellow Bellflower’ tree may have
families primarily for cooking, some of been the seed parent (Maas, 1970a;
which are too acidic for eating fresh, Khanizadeh and Cousineau, 1998), or
declined as populations became more ‘Delicious’ may have arisen as a sprout from
urban and lifestyles changed. Many older a seed or a seedling rootstock. Mr Hiatt was
cultivars have specific flaws that prevent impressed by the fruit and named the culti-
commercial production, or their appear- var ‘Hawkeye’. It was eventually purchased
ance deters a majority of consumers, but by Stark Brothers Nursery, who renamed it
they may be fine for home gardens where ‘Delicious’ and introduced it commercially in
individual flavour preference and dis- 1895. In its various forms, ‘Delicious’ has
ease/pest resistance are the criteria of become the world’s most important and
prime importance. best-studied cultivar. It has long been the
French (1970) gives an interesting backbone of the US industry. With its high
account of old American cultivars and why colour and distinctive appearance,
they were displaced. Specific faults respon- ‘Delicious’ is frequently used as a generic
sible for the decline of these old cultivars apple in advertisements and artwork.
include: lack of market demand or inability ‘Delicious’ is an important cultivar in the
to compete with newer cultivars; pro- USA (especially Washington), the European
nounced biennial bearing; inadequate Union, Australia, China and many other
winter-hardiness; fatal disease suscepti- countries. Synonyms: ‘Hawkeye’, ‘Red
bility; narrow adaptation; low yield; lack of Delicious’, ‘Stark Delicious’.
precocity; severe preharvest drop; severe
water-core; mediocre eating quality; in- 4.2.1.1 Tree
sufficient fruit size; unattractive fruit; poor
Standard trees moderately vigorous,
storage ability.
upright-spreading, spurring fairly freely,
basitonic, Lespinasse type II. Medium in pre-
cocity, productivity and regularity of bear-
4.2 Cultivar Descriptions ing. Widely adaptable, but performs best in
areas with warm summers, high light inten-
World production of apples is covered in sity and adequate water-supply. Bears pri-
Chapter 2 of this volume. Here we profile the marily on spurs. Much of modern
12 apples that are currently the most impor- production is on spur-type trees, i.e. trees
tant (by tonnage) in world trade. Tree and with a high proportion of the laterals being
fruit descriptions are presented in note form. spurs instead of long shoots (classic type I in
Selected pest-susceptibility reports are cited, the Lespinasse system). Spur types tend to
but should be interpreted with caution. Pest be more upright and require limb spreaders.
resistance is considerably influenced by cli- They are usually more precocious than stan-
mate and cultural practices as well as the dard strains and a little smaller in final tree
genetics of the host and pest. Detailed size. Diploid, not self-fertile; blooms mid-
descriptions of the bacteria and fungi season. Moderately to very cold-hardy
(Chapter 18) and insects and arachnids (Maas, 1970a; Kabluchko and Grigorenko,
(Chapter 19) mentioned in this chapter are 1976; Khanizadeh and Cousineau, 1998);
covered elsewhere in this volume. hardy to US Department of Agriculture
Characteristics of Commercial Apple Cultivars 63
(USDA) zone 4b (−34°C) (Strang and 4.2.1.3 Storage and postharvest

Stushnoff, 1975; Luby et al., 1999). King
‘Delicious’ stores 3–4 months in air (Smock
blooms more sensitive to frost than those of
and Neubert, 1950). Optimal controlled-
‘McIntosh’ or ‘Golden Delicious’, similar to
atmosphere (CA) storage recommendations
‘Jonagold’, and less sensitive than ‘Fuji’
vary with region, ranging from 0.7 to 2.5%
(Marro and Deveronico, 1979; Shibata and
O2, 0 to 4.5% CO2, −0.5 to 1.1°C, for storage
Mizuno, 1988; Mittelstadt, 1989; Mittelstadt
of 6–11 months (Kupferman, 1997).
and Salzer, 1989). Chilling requirement
‘Delicious’ is not chilling-sensitive or sensi-
medium to high, about 600–800 h in some
tive to low O2. It is prone to scald, particu-
estimations (Semadi, 1988) and 1200–1300
units in others (Ghariani and Stebbins, 1994). larly if picked too early (125–135 DAFB).
Resistant to heat, drought and hail (Khalin, Spur types should be picked 7–10 days later
1971; Timoshenko, 1976; Khalin and Dzetsin, than standard strains for comparable scald
1980). Not prone to preharvest drop. control (Fisher and Ketchie, 1989). The lobes
are sensitive to heat injury and can develop
symptoms resembling scald (Meheriuk and
4.2.1.2 Fruit McPhee, 1984).
See Bultitude (1983). Harvested at 140–160 ‘Delicious’ is also susceptible to water-
days after full bloom (DAFB) (Smock and core, especially as maturity advances. Later-
Neubert, 1950; Maas, 1970a; Khanizadeh and harvested fruit can also become mealy in
Cousineau, 1998) in a single pick. Maturity storage. ‘Delicious’ can get bitter pit, but it is
indices include DAFB, ground colour change preventable by proper fertilization practices
from green towards yellow, flesh colour and crop-load management. It is susceptible
change from green to cream, starch index to mouldy core.
and soluble solids. Size medium to large;
adequate thinning important for fruit size 4.2.1.4 Production notes
and return bloom. Shape oblong conic to
truncate conic, sometimes waisted below ‘Delicious’ is susceptible to apple scab, resis-
apex; prominently ribbed and irregular with tant to powdery mildew and highly resistant
very pronounced crowning at apex (Plate to fire blight. It is resistant or slightly suscep-
4.1). Stem cavity wide and fairly deep, usu- tible to cedar-apple rust, and susceptible to
ally russet-free. Stems medium, stout, some- quince rust and hawthorn rust (Aldwinckle,
times fleshy, exserted. Calyx basin wide and 1974; Warner, 1992). ‘Delicious’ is also mod-
deep, distinctly ribbed; eye medium, a little erately susceptible to Botryosphaeria canker,
open. Skin very tough, resistant to bruising, very susceptible to Valsa canker (Bessho et
russet-free, dry, smooth and glossy. Ground al., 1994) and susceptible to Nectria canker,
colour greenish yellow, overcolour varies black-rot canker (Miller, 1973), Monilinia
with strain; original said to be strawberry fructigena but not Monilinia laxa (Massodi
red with darker stripes (Maas, 1970a); and Bhat, 1988; Reznikova, 1990),
newer strains can be close to 100% dark Clathridium corticola fruit rot (Thind et al.,
crimson. Flesh cream, sometimes tinged 1975), leaf spot (Macek, 1974), Alternaria leaf
with green, very firm, fine-grained, juicy, blotch (Saito and Takeda, 1984) and apple
mealy if overripe. Flavour sweet, low in scar skin viroid (Desvignes et al., 1998). It is
acid, aromatic, distinctive. resistant to crown rot (Aldwinckle et al.,
1975) and brown leaf spot (Koropatyuk,
USES Very good for dessert, fair to poor for 1974) and tolerant of apple mosaic virus
culinary uses, fair for sauce, good for juice, (Singh et al., 1981).
fair to poor for drying, good for minimally ‘Delicious’ is very prone to red-mite
processed slices (Smock and Neubert, 1950; infestation (Smock and Neubert, 1950;
Kim et al., 1993b; Root, 1996; Lisowa et al., Fisher and Ketchie, 1989; Khanizadeh and
1997). Exceptionally good for handling and Cousineau, 1998), but is frequently consid-
shipping. ered tolerant to damage from this pest
(Beers and Hull, 1987). ‘Delicious’ is suscep- 1919. Earlier colouring was important to
tible or very susceptible to mullein bug growers, because the original strain often
(Campylomma verbasci Meyer) (Thistlewood coloured so late in maturation that the fruit
et al., 1989), Operophtera brumata Linnaeus, would become mealy in storage. The name
Rhopsalosiphum insertum Walker and rosy ‘Starking Delicious’ has been given to a
apple aphid (Cottwald, 1987). It was found succession of superior standard strains by
to be more resistant than some cultivars to Stark Brothers Nursery, beginning with the
maize weevil (Lorenzato and Grellmann, colour sport found by L. Mood in 1921.
1987), tufted apple bud moth (Knight and Most spur types have been whole-tree
Hull, 1988), apple rust mite (Bulgak, 1981), mutations (Fisher and Ketchie, 1989). In the
two-spotted mite (Yiem, 1993), woolly apple mid-1950s, ‘supercolour’ sports began to
aphid (Asante, 1994) and apple maggot emerge, such as the spur-type ‘Starkrimson
(Goonewardene et al., 1979). Delicious’ strain found by R. Bisbee in 1951
‘Delicious’ is not very sensitive to sun- (Maas, 1970a). Sports include: ‘Starking
burn. Problems may occur regionally with Delicious’, ‘Starkrimson Delicious’, ‘Royal
internal bark necrosis, bud union necrosis, Red’, ‘Morgan Spur®’, ‘Oregon Spur® II’,
poor fruit set, variable shape and dead spur ‘Midnight™ Red Spur Delicious’, ‘Apex’,
disorder (Fisher and Ketchie, 1989; Fear and ‘Topred™’, ‘Imperial Double Red
Domoto, 1998). Delicious’, ‘Ace® Spur Red Delicious’,
‘Ultrared’, ‘Silverspur’, ‘Hardispur
Delicious’, ‘Sturdeespur Delicious’, ‘Red
4.2.1.5 Breeding
Zenith® Spur’, ‘Scarlet Spur®’, ‘Redchief®’,
‘Delicious’ is one of the top five most fre- ‘Super Chief®’ and many others.
quent founding clones of modern apple cul-
tivars, and has been especially used in
breeding in the USA and Pacific rim (Noiton 4.2.2 ‘Golden Delicious’
and Alspach, 1996). It is a source of resis-
tance to fire blight and powdery mildew ‘Mullins Yellow Seedling’ was a chance
and of long storage life. Cultivars with seedling found by A.H. Mullins of Clay
‘Delicious’ in their parentage include ‘Fuji’,
County, West Virginia, USA. The date of
‘Gala’, ‘Empire’, ‘Melrose’, ‘Earlidel’,
origin is either 1890 (Smith, 1971; Bultitude,
‘Sekaiichi’, ‘Huashuai’, ‘Jinguang’, ‘Jupiter’,
1983) or 1905 (Percival and Proctor, 1994;
‘Summerdel’ and the scab-resistant culti-
Baugher and Blizzard, 1998). In 1914, prop-
vars ‘Priscilla’, ‘Enterprise’, ‘Florina’ and
agation rights were sold to Stark Brothers
‘McShay’, among others.
Nursery, who introduced the cultivar as
‘Golden Delicious’ (Baugher and Blizzard,
4.2.1.6 Sports 1998). The parentage of ‘Golden Delicious’
Over 100 strains of ‘Delicious’ exist; the is unknown, but is speculated to be either
original strain is scarcely grown at present. ‘Grimes Golden’ open pollinated (Smith,
All strains offer improved colour (timing 1971; Bultitude, 1983) or ‘Grimes Golden’ ×
and/or amount, in stripe or blush patterns) ‘Golden Reinette’ (Maas, 1970b;
and many are also spur types. Strains vary Khanizadeh and Cousineau, 1998). ‘Golden
in many other traits, including productiv- Delicious’ is undoubtedly the most impor-
ity, alternate bearing, fruit drop, suscepti- tant yellow apple in the world, and is sec-
bility to disorders, fruit shape, maturity ond only to ‘Delicious’ in world
time, fruit set, flesh greenness, sugar and production. ‘Golden Delicious’ is grown
acid content, firmness, date of full bloom, very widely, but is especially popular in
winter-hardiness and consumer acceptance. Europe and the USA. It has, however, lost
Most strains originated as spontaneous ground to newer cultivars such as
mutations in commercial orchards. The first ‘Jonagold’, ‘Elstar’, ‘Gala’ and ‘Pink Lady®’
colour sport was ‘Richared Delicious’ in in some areas.
4.2.2.1 Tree stripes. Lenticels conspicuous, fairly large,

grey-brown dots, prone to russet. Flesh
Moderately vigorous, spreading, spurs very
cream, firm, crisp, tender, fine-grained, juicy.
freely, wide branch angles, mesotonic. Very
Flavour sweet, a little acid, aromatic.
easy to manage, adaptable to a wide range of
Oxidation of cut flesh is fairly slow (Root,
soil and climatic conditions. Fruit finish and
1996; Khanizadeh and Cousineau, 1998).
texture best in warm areas with dry sum-
mers (Maas, 1970b). Bears on 1-year-old USES Very good for dessert and culinary
wood, terminals and spurs (type III in the use. Maintenance of flesh integrity makes it
Lespinasse system). Precocious, usually suitable for slices (Root, 1996), including
annual-bearing, highly prolific, not subject to minimally processed slices (Kim et al.,
preharvest drop. Diploid with abundant 1993b). Good for sauce, dehydration and
pollen, slightly self-fertile (Maas, 1970b; baby food and fair for juice (Maas, 1970b;
Pasqual et al., 1981; Niu et al., 1994). Blooms Root, 1996; Lisowa et al., 1997).
mid-season. Usually considered only moder-
ately cold-hardy. Reports disagree on its sen-
sitivity to bloom frost (high: Nybom, 1992; 4.2.2.3 Storage and postharvest
low: Mittelstadt and Koch, 1979; Mittelstadt, ‘Golden Delicious’ stores 3–4 months in air
1989), but it appears to be more resistant (Smock and Neubert, 1950). Optimal CA con-
than ‘Delicious’. Chilling requirement ditions vary with region, ranging from 1.0 to
reported as 600–800 h (Tabuenca and 3.0% O2, 0 to 4.0% CO2, −0.5 to 2.0°C, for
Jimenez, 1984; Semadi, 1988; Barahona et al., 6–10 months of storage (Kupferman, 1997).
1992) or 1275 units (Ghariani and Stebbins, ‘Golden Delicious’ is sensitive to bruising
1994). Considered fairly drought-resistant in from rough handling or mechanical impact,
Turkmenistan (Kosheleva et al., 1986). Flood and bruised fruit is susceptible to storage
tolerance intermediate, less than ‘Jonathan’ rots. Fruit is usually waxed to prevent skin
(Lee et al., 1983). shrivelling in storage. Texture becomes
mealy if harvested too late (Watada and
Abbott, 1985). Susceptibility to scald varies
4.2.2.2 Fruit
with growing conditions; it can be prevented
See Smith (1971), Bultitude (1983), with diphenylamine (DPA), ethoxyquin or
Khanizadeh and Cousineau (1998). low-O2 CA storage (Kupferman, 1997). In
Harvested 135–150 DAFB (Smock and some areas DPA is not recommended as it
Neubert, 1950; Maas, 1970b; Khanizadeh and causes a blue-grey skin discoloration and
Cousineau, 1998), usually in a single pick, ‘fixes’ chlorophyll in the skin (Lau, 1986).
when the ground colour starts to change Ultra-ultra-low oxygen (UULO) or pre-
from green to yellow (sometimes earlier for treatment with 15% CO2 prior to CA
CA storage). Size medium to large; round- improves firmness retention at 0°C (Watada
conic to oblong, ribbed on body and dis- and Abbott, 1985; Resnizky and Sive, 1991).
tinctly five-crowned at apex (Plate 4.2). Stem
cavity deep and rather narrow, with some
4.2.2.4 Production notes
russet. Stems long, slender, well exserted.
Calyx basin medium in depth and width, ‘Golden Delicious’ is very susceptible to
ribbed, sometimes partly russeted. Eye apple scab. Susceptibility to powdery
closed or partly open. Skin dry, tender, thin, mildew appears to vary among regions, with
smooth and prone to russet in certain cli- reports of both high and low susceptibility
mates, especially if humid. Russet-resistant being common. ‘Golden Delicious’ is suscep-
sports and gibberellins A4 + A7 (GA4+7) tible to quince rust and certain races of
sprays sometimes used commercially to cedar-apple rust (Aldwinckle, 1974; Chen
reduce skin russet. Ground colour dull and Korban, 1987; Warner, 1992). It is usually
greenish yellow, becoming golden yellow, rated as moderately resistant to fire blight.
sometimes with a slight orange flush but no ‘Golden Delicious’ is among the few culti-
vars with partial resistance to Nectria canker for high fruit quality, precocity and good tree
(Grabowski, 1987, 1994; van de Weg, 1989). It habit. In Germany, ‘Golden Delicious’ prog-
is partially resistant to Alternaria blotch eny were noted as having a high percentage
(more than ‘Delicious’, less than ‘Jonathan’) of seedlings with good resistance to fire
(Sawamura, 1972; Bulajic et al., 1996). It is blight (Fischer et al., 1996). ‘Golden Delicious’
resistant to M. laxa Aderhold & Ruhland is one of the top five most frequent founding
Honey (Reznikova, 1990) and partially resis- clones of modern apple cultivars (Noiton and
tant to Pezicula malicortis H. Jacks. Nannf. Alspach, 1996). Its progeny are poised to
and silver leaf (Borecki and Czynczyk, 1985) dominate world production in this century
and to black-rot canker (Miller, 1973). It is (Percival and Proctor, 1994). Apples with
more resistant than ‘Delicious’ to ‘Golden Delicious’ in their parentage include
Botryosphaeria canker (Latorre and Toledo, ‘Mutsu’ (‘Crispin’), ‘Jonagold’, ‘Elstar’,
1984). ‘Golden Delicious’ is susceptible to ‘Gala’, ‘Pink Lady®’ (‘Cripp’s Pink’),
fruit rot by Phytophthora cactorum (Lebert. & ‘Sundowner®’, ‘Corail’/’Pinova™’, ‘Arlet’
Cohn) Schroter (Mourichon and Salle, 1981; (‘Swiss Gourmet®’), ‘Delblush’ (‘Tentation®’),
Grove, 1993); it is susceptible to leaf blotch ‘Delcorf ’ (‘Delbarestivale®’), ‘Creston’,
(Janick et al., 1996) and Phoma leaf spot ‘Summerred’, ‘Fu Shuai’, ‘Sekaiichi’,
(Macek, 1974). ‘Golden Delicious’ is moder- ‘Shizuka’, ‘Pirella’, ‘Kinsei’, ‘Sweet Caroline’,
ately susceptible to apple mosaic virus ‘Falstaff’, ‘Xinguan’, ‘Tsugaru’, ‘Shamrock’,
and apple witches’-broom mycoplasma-like ‘Orin’, ‘Maigold’, ‘Delbard Jubilé’ and the
organism (Kegler et al., 1992) and tolerant of scab-resistant (Vf ) cultivars ‘Sir Prize’,
apple scar skin viroid (Desvignes et al., 1998). ‘GoldRush’, ‘Primiera’, ‘Ariwa®’, ‘Florina’,
‘Golden Delicious’ is preferred over ‘Princesa’, ‘Enterprise’, ‘Baujade’,
‘Cox’s Orange Pippin’ by egg-laying ‘Freedom®’, ‘Priam’ and ‘Prima’.
codling moths (Blago and Dickler, 1990) and
less preferred than ‘Stayman’ or ‘Rome
4.2.2.6. Sports
Beauty’ by feeding larvae of tufted apple-
bud moth (Knight and Hull, 1988; Meagher Numerous strains exist, varying in russet
and Hull, 1991). It is susceptible to apple resistance, spurriness, blush, fruit size, earli-
fruit moth in India (Argyresthia conjugella ness of bearing, tree size, yield efficiency,
Zeller) (Khajuria et al., 1987), susceptible to productivity, alternate bearing and powdery-
red mite (Kolbe, 1972; Skorupska, 1992), mildew resistance. Götz and Silbereisen
more resistant than ‘Fuji’ or ‘Delicious’ to (1989) list over 60, including ‘Starkspur
two-spotted mite (Yiem, 1993; Yiem et al., Golden Delicious’, ‘Smoothee®’ (‘Gibson
1993), less resistant than ‘Delicious’ to apple Golden Delicious’, ‘Golden Glory™’,
maggot (Goonewardene et al., 1979) and ‘Goldspur® Delicious’), ‘Lutz’, ‘Lysgolden’,
highly susceptible to rosy apple aphid (Graf ‘Belgolden’ and ‘Reinders’. None exceeds
et al., 1992). standard ‘Golden Delicious’ in acceptance
‘Golden Delicious’ can develop bitter pit (Percival and Proctor, 1994).
or sunburn under appropriate conditions,
but is not considered highly prone to these
disorders. Calyx green-end disorder, a flat- 4.2.3 ‘Fuji’
tening and distortion of the calyx end with
persistent green colour of the skin, appears ‘Fuji’ is the offspring of a ‘Ralls Janet’ ×
to be a result of fluoride toxicity (Seeley, ‘Delicious’ cross made in 1939 (Smith, 1971;
1979; Barritt and Kammereck, 1983). Kikuchi et al., 1997). It was named ‘Fuji’ in
1962 by the Horticulture Research Station
in Morioka, Japan. The name commemo-
4.2.2.5 Breeding
rates Fujisaki in Aomori, Japan, where the
‘Golden Delicious’ has been used extensively cross was made (Yoshida et al., 1998). ‘Fuji’
in breeding, especially in the Pacific rim, USA is the most important apple in Japan and
and western Europe. It is valued as a source China and is a major cultivar in Korea,
Brazil, Argentina, Chile and Australia. It without special management. Elaborate

has been planted extensively in both hemi- practices to boost colour development have
spheres in the past decade. About 80% of been developed in Japan, including bagging
current ‘Fuji’ acreage is located in China (Arakawa, 1998), leaf removal, reflective
(Avermaete, 1999). ‘Fuji’ has a long storage mulches and fruit turning. Flesh cream,
and shelf life, perhaps because of its low above average in crispness, firmness, juici-
ethylene production and low respiration ness, fine-grained. Flavour sweet and very
rate (Yoshida et al., 1998). mild, high in sugar and low in acidity. Low
in dietary fibre compared with other culti-
vars (Gheyas et al., 1997).
4.2.3.1 Tree
Vigorous, spreading and willowy, mesotonic, USES Excellent for dessert, good for process-
precocious, productive but somewhat slow ing quality, except cider (Yiem et al., 1980).
to spur in initial years (Tustin, 1994) and pro-
duces some blind wood. Tip bearer
4.2.3.3 Storage and postharvest
(Lespinasse type IV), with long darts; flow-
ers on 1- and 2-year-old wood. Difficult to ‘Fuji’ is highly prone to water-core, prone to
thin chemically, strong tendency to bear stem-end cracking, bitter pit, cork spot and
biennially. No preharvest drop. Diploid, external brown staining. Severe water-core
requires a pollinizer, blooms in mid-season. shortens storage life and may cause internal
Flooding tolerance intermediate (Lee et al., breakdown. Compared with other cultivars,
1983). Trees winter-hardy but with a lower ‘Fuji’ has a very slow rate of firmness loss
chilling requirement than ‘McIntosh’ or and a long shelf life (Yoshida et al., 1998).
‘Delicious’ (Ghariani and Stebbins, 1994); For short-term storage, CA is no better than
estimations of chilling requirement vary regular atmosphere storage for firmness
from 600–800 h (Barahona et al., 1992) to 1050 retention, but CA improves retention of
units (Ghariani and Stebbins, 1994). More acidity (Drake, 1993). ‘Fuji’ is susceptible to
sensitive to bloom frost than ‘Delicious’ or internal browning in long-term CA (≥ 6
‘Jonagold’ (Shibata and Mizuno, 1988). months); the problem is worse with mature
fruit. Short-term (3 days) exposure of fruit
samples at harvest to 20 kPa CO2 may be
4.2.3.2 Fruit
useful as a predictor of susceptibility to
Harvested 140–180 DAFB, early to mid- internal browning (Volz et al., 1998). ‘Fuji’ is
November in Japan. May require several also susceptible to core browning, a disorder
picks, depending on colour management. whose incidence rises with later picking and
Ground colour, overcolour and starch disap- preharvest calcium treatments (Yoshida et
pearance are used as maturity indices, but al., 1998). Early-picked ‘Fuji’ may develop
not firmness or soluble solids (Argenta et al., scald, but it is preventable with DPA, ultra-
1995; Britz, 1998; Yoshida et al., 1998). Size low oxygen (ULO) or hypobaric storage
large; shape round-conic to round-oblate, (Kupferman, 1997; Yoshida et al., 1998).
ribbed on body (Plate 4.3). Stem cavity wide Optimum CA conditions vary with region,
and moderately shallow, sometimes partly ranging from 0.7 to 2.5% O2, < 0.5 to 2.0%
lined with brown russet. Stems long, stout, CO2 at 0–1°C, for 7–11 months of storage
well exserted. Calyx basin wide, moderately (Kupferman, 1997).
deep, distinctly ribbed. Eye medium, closed,
anthers frequently persisting. Lenticels large
conspicuous white dots. Skin tough, smooth,
dull or sometimes roughened with russet. Bloom thinning is recommended, but some
Susceptible to stem punctures (Britz, 1998). chemicals cause russet (Jones et al., 1998;
Ground colour pale yellow-green with red Yoshida et al., 1998). Thinning to singles with
blush and darker stripes (Smith, 1971). 75 leaves per fruit is recommended in Japan
Colour frequently poor on standard strain (Yoshida et al., 1998).
‘Fuji’ is highly susceptible to apple scab 4.2.4 ‘Granny Smith’

and fire blight and moderately resistant to
powdery mildew. It is resistant to quince ‘Granny Smith’ is a chance seedling discov-
rust; cedar-apple rust lesions occurred on ered on the farm of Maria Ann and Thomas
leaves but not fruit in one 3-year study Smith of Ryde, New South Wales, Australia.
(Warner, 1992). ‘Fuji’ is considered suscepti- The original tree was fruiting by 1868.
ble or highly susceptible to Alternaria blotch ‘Granny Smith’ is believed to be an open-
(but less so than ‘Delicious’) (Brooks and pollinated seedling of ‘French Crab’, but it
Olmo, 1997; Khanizadeh and Cousineau, also resembles some of the American
1998; Yoshida et al., 1998), lenticel infection ‘Greening’ cultivars and ‘Cleopatra’
by apple rot (Botryosphaeria dothidea Moug. (Warrington, 1998). Although there have
Fr. Ces & DeNot) (Kim and Kim, 1989), fruit been significant plantings of ‘Granny Smith’
ring rot and twig cankers (Physalospora piri- since the 1920s in Australia, it has only
cola Dose) (Luo et al., 1996), Valsa canker become an important apple in world trade
(Bessho et al., 1994), bitter rot and Mucor rot since 1950. ‘Granny Smith’ is chiefly a south-
(Mucor piriformis E. Fisch) (Yoshida et al., ern hemisphere apple. It has been grown
1998). It is also sensitive to apple scar skin extensively in Australia, Argentina, Chile,
viroid (Desvignes et al., 1998) and the New Zealand and South Africa. Unmet con-
viruses russet ring and apple fruit crinkle sumer demand stimulated significant plant-
(Yoshida et al., 1998). ing in western North America and southern
‘Fuji’ is fairly resistant to apple maggot Europe in the 1970s. Today ‘Granny Smith’
(Lamb et al., 1988) and susceptible to two- accounts for a third of southern hemisphere
spotted spider mite (Yiem, 1993; Yiem et al., exports (Avermaete, 1999).
1993). ‘Fuji’ is very susceptible to fruit skin
or lenticel russeting and moderately prone to
4.2.4.1 Tree
sunburn (Britz, 1998).
Moderately vigorous, upright-spreading,
spurs fairly freely, but has some blind wood
4.2.3.5 Breeding
(Bultitude, 1983; Khanizadeh and
‘Fuji’ is valued as a parent for its large fruit, Cousineau, 1998). Precocious, bearing
excellent quality and long storage life. It is cur- heavy annual crops. Production of 120–130 t
rently being used in many breeding pro- ha−1 is routine in New Zealand, but yield is
grammes, especially on the Pacific rim. ‘Fuji’ frequently lower elsewhere (Warrington,
offspring include ‘Himekami’, ‘Hokuto’, 1998). In the Lespinasse system, ‘Granny
‘Huaguan’, ‘Huashuai’, ‘Senshu’ and the scab- Smith’ is type IV (acrotonic tip bearer). Fruit
resistant (Vf ) Brazilian cultivar ‘Catarina’. are initially borne on branch tips but later
also on laterals, primarily 1- and 2-year-old
wood. Partially self-fertile, diploid, partially
4.2.3.6 Sports
self-thinning (Warrington, 1998). Cold-har-
At least 128 named strains occur in Japan diness moderate, adequate for central
alone (Komatsu, 1998). Colour sports are the Washington State but not Quebec or
most common. Strains vary in stability, Minnesota (Khanizadeh and Cousineau,
colour pattern, intensity and amount of 1998; Luby et al., 1999). Blooms mid- to late
colour, ploidy level, spurriness, climatic season. High resistance to drought and
adaptation, eating quality and harvest time. moderate resistance to heat reported from
In Japan, the striped sports are highly the Ukraine (Khalin, 1989). Chilling require-
favoured, but blush sports may colour better ment reported as 400–600 h in Algeria (less
in warm climates (Komatsu, 1998). Other than ‘Delicious’ or ‘Golden Delicious’),
mutants include ‘Beni-Shogun’, ‘Seirin 600–800 h in Ecuador, 1040 units in the
Spur’, ‘Tensei’ (tetraploid), ‘Sun Fuji™’, USA, similar to ‘Fuji’ or ‘Jonagold’ (Semadi,
‘Myra Red Fuji’, ‘Jubilee Fuji’, ‘Yataka’ and 1988; Barahona et al., 1992; Ghariani and
‘Takano Wase’. Stebbins, 1994).
4.2.4.2 Fruit mildew and very susceptible to fire blight

(Jeger et al., 1986; Khanizadeh and
Harvested 170–210 DAFB in a single pick
Cousineau, 1998), but is resistant to cedar-
(Janick et al., 1996; Khanizadeh and
apple rust and quince rust (Warner, 1992;
Cousineau, 1998; Warrington, 1998). Size
Khanizadeh and Cousineau, 1998). It is more
medium to large; shape round to round-
resistant than ‘Delicious’ to Botryosphaeria
conic, fairly regular, slightly flattened at base
trunk canker (Latorre and Toledo, 1984) and
and apex, slightly five-crowned at apex, uni-
is tolerant of apple scar skin viroid
form (Plate 4.4). Stem cavity fairly narrow
(Desvignes et al., 1998). Compared with
and deep, sometimes partly lined with grey
‘Delicious’ it is more resistant to red mite
russet. Stems moderately long and slender, (Monetti and Fernandez, 1996) but less resis-
level with base or exserted. Calyx basin tant to woolly apple aphid (Asante, 1994).
medium in width and depth, distinctly
ribbed, puckered and russet-free. Eye is
medium, closed or slightly open. Skin thick, 4.2.4.5 Breeding
tough, resists bruising, smooth, waxy, becom- ‘Granny Smith’ is a parent of the French cul-
ing greasy with maturity, lacks bloom. tivars ‘Baujade’ (Vf ) and ‘Delgaly’.
Ground colour bright green, becoming green-
ish yellow, with large conspicuous areolar
white lenticels. Sometimes a slight pink blush 4.2.4.6 Sports
present but no stripes. Flesh greenish white, Sports include ‘Granspur’, ‘Greenspur’,
very firm, rather coarse, juicy, subacid, tart- ‘Earlee Grannee’. These strains reportedly dif-
sweet, refreshing, but lacking in flavour. fer from the standard in fruit size, flesh colour
(whiter), tree spurriness or compactness,
USES Good dual-purpose dessert and culi- ripening time, fruit set, internode length, lenti-
nary apple. Fruit especially high in pectin cel prominence, productivity and the presence
(Blagov, 1998). of dark green stripes in ‘Granspur’ (Brooks
and Olmo, 1997). Most plantings are still stan-
4.2.4.3 Storage and postharvest dard types. Some believe that spur types have
poorer fruit quality (Warrington, 1998).
‘Granny Smith’ is a long-keeping apple, pos-
sibly because of its low ethylene production
(Warrington, 1998). It is very susceptible to 4.2.5 ‘Gala’
superficial scald, particularly if picked too
early, but this disorder can be controlled ‘Gala’ is the product of two generations of
with DPA and/or ULO storage. Premium controlled crossing by amateur breeder J.H.
postharvest quality is attained when the fruit Kidd of Wairarapa, New Zealand (White,
is picked with a medium green ground 1998). ‘Gala’ is the offspring of a cross of
colour and when all starch has disappeared ‘Kidd’s Orange Red’ × ‘Golden Delicious’
from the core area (Warrington, 1998). made about 1934. ‘Kidd’s Orange Red’ was
Optimal CA conditions range from 0.8 to itself a cross of ‘Delicious’ × ‘Cox’s Orange
2.5% O2, 0 to 5.0% CO2, −0.5 to 2°C, for 7–11 Pippin’ (Noiton and Alspach, 1996). The
months of storage (Kupferman, 1997). apple was named ‘Gala’ in 1962, and
Incidences of core flush and scald are higher released for commercial planting in 1965.
in warm climates. ‘Granny Smith’ may However, it did not become really popular
develop bitter pit and water-core if picked until the mid-1970s when several red colour
too late (Khanizadeh and Cousineau, 1998). sports appeared (Tustin, 1990). ‘Gala’ is an
important cultivar in New Zealand, Brazil,
Argentina, Chile, Australia, China, the USA
and Europe (especially France), and has been
‘Granny Smith’ is susceptible to scab, moder- planted extensively in both hemispheres in
ately to highly susceptible to powdery the past decade. Synonym: ‘Kidd’s D.8’.
4.2.5.1 Tree bright orange-red and strewn with deeper

red stripes. Lenticels fairly inconspicuous.
Moderately vigorous, upright-spreading,
Flesh pale yellow, juicy, firm, crisp, fine-
with long pliant branches, heavily spurred,
grained. Flavour sweet, low in acid, refresh-
mesotonic habit. Type III in the Lespinasse
ing, aromatic, excellent quality.
system, like ‘Golden Delicious’ but with nar-
rower branch angles. Fruit borne on 1- and 2- USES Main use is dessert. Suitable for dry-
year-old wood, terminals and long strong ing (Lisowa et al., 1997).
bourse shoots (Tustin, 1990). Highly preco-
cious, bears prolific annual crops.
Devigorating burr knots may form at the 4.2.5.3 Storage and postharvest
base of older limbs (Tustin, 1990). Wood is ‘Gala’ has no significant storage disorders
brittle and prone to breakage. Easy to train but only a medium-term storage life (Tustin,
with modern methods, widely adapted. Fruit 1990). CA recommendations vary with
size and colour very responsive to light region and strain; optimum conditions range
exposure. Cold-hardiness moderate, similar from 1 to 3% O2, < 0.5 to 2% CO2, 0 to 3°C for
to ‘Golden Delicious’ (Luby et al., 1999). storage of 4–9 months (Kupferman, 1997).
Chilling requirement estimates vary from
600–800 h (Barahona et al., 1992) in Ecuador
to 1150 units in Oregon, USA (Ghariani and 4.2.5.4 Production notes
Stebbins, 1994). Diploid, somewhat self-fer- ‘Gala’ is highly susceptible to apple scab
tile. Extended bloom, starting mid-season. (especially on fruit) and fire blight, suscepti-
Fruit set heavy; easy to thin chemically and ble to Nectria canker and silver leaf and
not prone to biennial bearing. Fruit quality moderately susceptible to powdery mildew.
best on 2-year-old spurs; renewal pruning is It is moderately susceptible to cedar-apple
recommended (Tustin, 1990). rust and resistant to quince rust (Warner,
1992). It is highly resistant to Alternaria
blotch (Shin et al., 1986) and resistant to
4.2.5.2 Fruit
Botryosphaeria berengeriana deNot stem
See Smith (1971), Bultitude (1983), canker and fruit rot (Melzer and Berton,
Khanizadeh and Cousineau (1998). 1986). It is susceptible to russet ring virus,
Harvested 120–140 DAFB, 2–4 weeks before apple mosaic virus (Brooks and Olmo, 1997;
‘Delicious’. Fruit adhere well to tree and can Khanizadeh and Cousineau, 1998) and apple
be hard to pick; preharvest drop minimal. scar skin viroid (Desvignes et al., 1998).
Cull percentage typically low. Ripening ‘Gala’ is fairly resistant to maize weevil in
uneven; requires at least three picks, based Brazil (Lorenzato and Grellmann, 1987).
on ground colour change from green to
creamy yellow. Size small to medium and
4.2.5.5 Breeding
usually uniform; shape round-conic to
oblong-conic or oblong, with ribbing on ‘Gala’ is in current use in many breeding
body; distinctly five-crowned at apex (Plate programmes throughout the world. ‘Gala’
4.5). Stem cavity medium in width, deep to offspring include ‘Sansa’, ‘Chinook’, ‘Pacific
very deep, partly lined with grey russet. Rose’, ‘Scifresh’, ‘Pacific Beauty’, ‘Pacific
Prone to stem-end russet in some climates. Queen’ and the scab-resistant (Vf ) French cul-
Stems long and slender, exserted. Calyx tivar ‘Initial’. Many programmes have other
basin medium, eye closed or a little open. ‘Gala’ offspring under test.
Skin russet-free, smooth and glossy, becom-
ing greasy; not susceptible to bruising; occa-
4.2.5.6 Sports
sionally some scarf skin at base. Develops
stem-end cracking when fully to overmature. ‘Gala’ is prone to producing red colour
Ground colour creamy yellow to golden yel- sports, which vary considerably in stability.
low, partly to fully flushed and flecked with Both striped and blush sports exist; the tradi-
tional retail market is based on striped types. very downy. Blooms late mid-season, fertile
Strains include ‘Tenroy’ (‘Royal Gala®’), pollen, diploid, slightly self-fertile. Slight to
‘Imperial Gala™’ (‘Mondial Gala®’, medium susceptibility to bloom frost; tree
‘Mitchgla®’), ‘Regal Gala’, ‘Galaxy Gala™’, moderately winter-hardy. Suitable polliniz-
‘Scarlet Gala’, ‘Fulford Gala™’, ‘Brookfield® ers are listed in Kemp (1996).
Gala’, ‘Buckeye® Gala’, ‘Pacific® Gala’, ‘Gale
Gala™’, ‘Delaf Gala’, ‘Regal Prince’ (‘Gala
4.2.6.2 Fruit
Must®’) and ‘Waliser’ (‘Crimson™ Gala’).
Strains vary slightly in flower density, matu- Harvested 140–150 DAFB in multiple picks,
rity date, yield, fruit size and bruise suscepti- picks easily, no preharvest drop. Difficult to
bility in some trials. Usually internal quality get good fruit size (Rom, 1998). Suitable for
does not vary (Kappel et al., 1992; Greene warm climates; in cooler areas, poorer eating
and Autio, 1993). quality (less aromatic). However, fruit from
cooler sites stores better. Moderately crisp,
quite juicy, sweet, refreshingly subacid to
4.2.6 ‘Jonathan’ tart, but, on unsuitable sites, a woody metal-
lic taste can occur. Flesh fairly firm, fine-tex-
‘Jonathan’ is believed to be a seedling of tured, white, slightly green to yellow, with
‘Esopus Spitzenburg’ (or ‘Spitzenberg’). In weak, characteristic aroma. Only moderately
1826, it was described by Judge Buel of tasty. Attractive, bright (sometimes pale or
Albany, New York State, USA, who named it brownish), solid crimson-red blush, with
after Jonathan Hasbrouck, the man who some short, broken red stripes, on a yellow-
drew his attention to the tree on the farm of green ground. Skin with finely netted, some-
Philip Rick in Ulster County, New York State. times patchy russet; slightly waxy, smooth,
‘Jonathan’ is still an important cultivar in the dry, lustrous with slight bloom and skin
USA (Michigan), several eastern European hammering; conspicuous, tiny lenticels. Size
countries and Japan. In western Europe, it medium to rather small, shape oblong to
has been superseded by more recent culti- round-conical, usually regular and symmet-
vars, such as ‘Jonagold’, but plantings are rical; fruit slightly ribbed, mainly at apex
still present in Switzerland, Korea, Austria, and in basin (Plate 4.6). Can be flat-sided.
Italy, Germany, New Zealand and Australia. Basin quite narrow, deep, often puckered,
Several good descriptions of ‘Jonathan’ are with quite small, closed eye; usually russet-
available, e.g. Baldini and Sansavini (1967), free. Stem cavity fairly narrow, deep, usually
Nilsson (1987), Sanders (1988), Götz and with some russeting, sometimes streaking
Silbereisen (1989), Friedrich and Petzold over shoulder. Stalk rather slender, short to
(1993), Morgan and Richards (1993), long. Skin thin to moderately thick, tough,
Manhart (1995), Khanizadeh and Cousineau hard, chewy.
(1998) and Rom (1998). Synonyms: ‘New
(Esopus) Spitzenberg’, ‘Philip(p) Rick’, USES Suitable for processing, cider, dessert,
‘Ulster (Seedling)’, ‘Johnathan’, ‘King sauce, juice and pies. Shelf life very good to
Philip(p)’, ‘Pomme Jonathan’, ‘Djonathan’, good.
‘Dzhonatan’.
4.2.6.1 Tree
‘Jonathan’ stores for 5 months in air, starting
Moderately vigorous to rather weak, good at 3–4°C, later 0–1°C, or for 6 months in
central leader, rather dense, feathered, ULO or in CA at 0–1°C, 1–5% CO2 and
spreading to weeping; wood twiggy, rather 1.5–3% O2 (Kupferman, 1997). Storage above
thin. Precocious, regular bearing, fairly good 2–3°C prevents Jonathan spot, in which
productivity. Bears on 2- and 1-year-old small black areas surrounding the lenticels
wood and somewhat on spurs, type III in the expand and sometimes become infected by
Lespinasse system. Leaves greyish green, secondary rots. CA storage prevents
Jonathan spot almost completely; in cold 4.2.6.5 Breeding

storage, spot can be severe, especially with
‘Jonagold’ is probably the best-known off-
ripe fruit. The onset of Jonathan spot is
spring of ‘Jonathan’, but many other culti-
delayed by rapid cooling in CA with high
vars are descended from it, including
CO2, but excess CO2 and low O2 damage
‘Idared’, ‘Jonadel’, ‘Melrose’, ‘Megumi’,
fruit. Storage in ULO, 1.5% O2 and 1.0% CO2
‘Monroe’, ‘Jonalicious’, ‘Akane’, ‘Undine’,
reduces brown core. Large fruit are prone to
‘Herma’, ‘Eva’, ‘Bonza’, ‘Rubinovoe Duki’,
decay. Fruit with water-core are not suitable
‘Malling Kent’, ‘Blushing Golden’, ‘Goldjon’,
for CA. ‘Jonathan’ is susceptible to scald,
‘Cadel’, ‘Cacanska Pozna’ and ‘Septer’. See
bruising, water-core, internal browning,
also Smith (1971) and Rom (1988). ‘Fiesta’
Jonathan spot; it has medium susceptibility
(‘Red Pippin®’) and ‘Elise’ (‘Roblos®’) are
to bitter pit, cork spot and low-temperature
both F2 offspring. It has also been used as a
browning (LTB). No shrivelling. The risk of
parent in the Ukraine. ‘Jonathan’ is one of
internal browning and spot increases with
the top five most frequent founding clones of
long storage. Jonathan breakdown (under
modern apples (Noiton and Alspach, 1996).
the skin) and senescent breakdown (from
core) can occur.
4.2.6.6 Sports
4.2.6.4 Production notes A number of sports have been identified,
Deep, light soils are suitable. Night frosts on mainly for colour. Examples: ‘Anderson
fruit just before harvest cause internal Jonathan’, ‘Kap(p)ai Red Jonathan’,
browning. Colour is often poor on standard ‘Jonared’, ‘Blackjon’ (‘Red Jonathan’), ‘Rode
‘Jonathan’. Summer pruning can improve Jonathan Fleuren’, ‘Rode Jonathan Heines’,
fruit colour. Repeated selective picking can ‘Watson Jonathan’, ‘Ruby Jon®’, ‘Jonnee’.
reduce storage problems. Incidence of Some tetraploids and some irradiated, spur-
Jonathan spot and internal browning type mutants are known (Götz and
increases with late picking and larger fruit Silbereisen, 1989).
size. Timely picking is necessary for good
internal storage quality. ‘Jonathan’ can pro-
duce parthenocarpic fruit after GA applica- 4.2.7 ‘Jonagold’
tion (Thomas, 1963). Early thinning is
required; it has a slight tendency to biennial ‘Jonagold’ resulted from a ‘Golden Delicious’
bearing. If trees are biennial or too vigorous, × ‘Jonathan’ cross made in 1943 at the New
root pruning in combination with supple- York State Agricultural Experiment Station
mental irrigation is a useful tool (Ferree, breeding programme in Geneva, New York
1992). Ethephon, naphthaleneacetic acid State, USA, and was introduced in 1968. It
(NAA) and carbaryl are effective thinning was named after its parents. ‘Jonagold’ is
agents (Windle and van Dam, 1989). Calcium highly appreciated by consumers for its eat-
sprays, dips or infiltration can reduce bitter ing quality. It has found special favour in
pit, cork spot, breakdown (Dewey et al., 1982; Belgium, where it accounts for ~60% of pro-
Fukuda, 1984) and Alternaria rot (Byun and duction (Lambrechts, 1994). ‘Jonagold’ is also
Chang, 1991). an important cultivar in The Netherlands,
‘Jonathan’ is very susceptible to mildew Germany, France, Switzerland, Italy, the UK,
(Janse et al., 1998), slightly to moderately sus- Japan, Australia, China, the USA and
ceptible to scab (Rati et al., 1994) and suscep- Canada (Way and Brown, 1998).
tible to cedar-apple rust (Joung et al., 1987),
fire blight and bitter rot (Rom, 1998); it is
4.2.7.1 Tree
resistant to Alternaria leaf spot and has strik-
ingly low susceptibility to Nectria canker Vigorous, spreading, large, with good
(Groenwold et al., 1994). branch angles and spurring freely. Type II in
the Lespinasse system, bears mainly on USES Very good for dessert, culinary use
spurs. Precocious, high-yielding, little pre- and processing (sauce, slices).
harvest drop (Way and Brown, 1998). Bears
annually in most districts, but can be bien- 4.2.7.3 Storage and postharvest
nial. Triploid; requires a pollinizer but can-
not donate pollen. Only moderately When picked at the optimum maturity,
cold-hardy, has suffered greatly in severe ‘Jonagold’ reportedly stores for up to 6
winters in Europe. Blossoms less sensitive to months in air at −0.5°C (Way and Brown,
bloom frost than ‘Fuji’, similar to ‘Delicious’, 1998); otherwise, it becomes soft and bland
and more sensitive than ‘Golden Delicious’, in long-term air storage. Commercial grow-
‘McIntosh’ or ‘Spartan’ (Shibata and ers may apply calcium three or more times to
Mizuno, 1988; Mittelstadt, 1989; Mittelstadt improve storage life and prevent bitter pit.
Maturity indices for long storage are given
and Salzer, 1989). Blooms in mid-season
by Lau (1992). Optimum CA conditions vary
with ‘Golden Delicious’. Chilling require-
among regions, ranging from 1.0 to 2.5% O2,
ment fairly high, ~1100 units (Ghariani and
1.0 to 4.5% CO2 at 0–3°C, for 5–10 months of
Stebbins, 1994). ‘Jonagold’ performs best in
storage (Kupferman, 1997).
cooler districts, such as northern Europe. In
hot areas, it suffers from sunburn, soft flesh
and poor colour. 4.2.7.4 Production notes
‘Jonagold’ is susceptible to scab, fire blight
and cedar-apple and quince rusts and highly
4.2.7.2 Fruit
susceptible to powdery mildew. It is suscep-
See Bultitude (1983), Götz and Silbereisen tible to Nectria canker, but less so than
(1989), Khanizadeh and Cousineau (1998), ‘McIntosh’ or ‘Gloster’ (Grabowski, 1994). In
Way and Brown (1998). Harvested 140–160 Korea it is considered highly resistant to
DAFB (Khanizadeh and Cousineau, 1998) Alternaria blotch (Shin et al., 1986). In certain
and requires three to four picks. Picking regions, it is subject to winter injury, sun-
date in New York State is just after burn, bitter pit, alternate bearing or excessive
‘Delicious’ (Way and Brown, 1998); earlier in size. Fruit colour can be poor, especially with
some other localities, with ‘Golden high crop load or strong vegetative growth.
Delicious’. Fruit large; round to round-conic
with very slight ribbing, moderately flat- 4.2.7.5 Breeding
tened and slightly five-crowned at apex
‘Jonagold’ is triploid and cannot be used in
(Plate 4.7). Stem cavity wide and fairly deep,
conventional breeding.
lined with grey russet. Stems fairly long and
moderately stout, exserted and often curved
to one side. Calyx basin fairly wide and 4.2.7.6 Sports
quite deep, somewhat ribbed. Eye small, a Around 100 strains of ‘Jonagold’ exist
little open. Skin smooth, prone to scarf skin (Goddrie, 1996), including ‘Jonagored
and frost blemishes, not prone to russet, (Morren®)’, ‘Schneica (Jonica®)’, ‘Jored’ (‘King
becoming greasy with maturity. Some sus- Jonagold’), ‘Jonaveld’ (First Red®), ‘Nicobel
ceptibility to bruising. Ground colour is Jonagold’, ‘Rubinstar® Jonagold’, ‘Decosta
bright yellow tinged with green, with short, Jonagold’, ‘Jonagold De Coster®’, ‘Jonabel
broad, broken, dull red stripes over 30–80% Jonagold’, ‘Novajo’, ‘Red Jonaprince (Red
of the surface. Lenticels small, grey and Prince®)’, ‘Marnica’, ‘Jonagold Boerekamp
green. Flesh light yellow, semi-firm, (Early Queen®)’. Most were selected for a
medium-grained, crisp, very juicy, slow to superior amount or intensity of red colour, but
brown after cutting. The flavour is subacid differences in fruit or tree shape, time of matu-
to sweet, aromatic, very rich. rity and yield have sometimes been reported.
4.2.8 ‘McIntosh’ 4.2.8.2 Fruit

See Beach et al. (1905b), Smock and Neubert
‘McIntosh’ is a chance seedling discovered in
(1950), Smith (1971) and Bultitude (1983).
1811 on the farm of John McIntosh near
Harvested 125–145 DAFB (about 4 weeks
Dundela, Ontario, Canada (Upshall, 1970).
Originally called ‘Granny’s apple’, it was before ‘Delicious’) in one or two picks. Size
renamed ‘McIntosh Red’ in 1836, later short- medium to large; round to oblate; size and
ened to ‘McIntosh’. Although ‘McIntosh’ shape uniform or slightly irregular, with
grafts were sold as early as 1835 (Upshall, slight to no ribbing (Plate 4.8). Stem cavity
1970), many references give 1870 as the date medium in width and depth, broadly fur-
of commercial introduction (Beach et al., rowed, acute to acuminate, partly lined
1905b; Smith, 1971; Bultitude, 1983; Proctor, with fine brown russet. Stems short, level
1998). The cultivar did not become popular with base, rarely exserted. Calyx basin
until around 1900, about the time that the medium in width and depth, smooth or
first scab sprays were developed. The slightly ribbed, sometimes with small fleshy
parentage of ‘McIntosh’ is a mystery, but its beads. Eye small, tightly closed to very
vegetative characteristics, cold-hardiness slightly open. Skin thick but tender, subject
and scab susceptibility have fuelled specula- to stem punctures, separates readily from
tion that the parents may include ‘Fameuse’ flesh, smooth, covered with conspicuous
(‘Snow Apple’), ‘Fall St Lawrence’ or the lilac bloom, glossy when buffed. Ground
Russian cultivar ‘Alexander’ (Upshall, 1970; colour green to yellow-green, 30–80%
Khanizadeh and Cousineau, 1998). After a deeply flushed with bright red (sometimes
severe winter in 1933/34 killed many dark, nearly purple overcolour) and faintly
‘Baldwin’ trees in the north-eastern USA, streaked with short broken carmine stripes.
they were replaced with the more cold-hardy Skin green where shaded. Flesh white,
‘McIntosh’ (Smock and Neubert, 1950). tinged with green or pink, sometimes
‘McIntosh’ is the leading cultivar in the veined with red, fine, very tender, juicy,
north-eastern USA and eastern Canada and firm but soon becoming soft. It is best
is important in eastern Europe. Synonym: adapted to cool areas with cold nights and
‘McIntosh Red’. clear autumn days. In warm areas, it has
poor colour, excessive preharvest drop and
4.2.8.1 Tree soft flesh. Sensitive to bruising.
Characteristic aromatic perfumed flavour,
Moderately vigorous, spreading, with good sprightly tart–sweet becoming nearly sweet.
branch angles, spurs freely, type III in the Very distinctive scent. Flesh browns quickly
Lespinasse system. Precocious, annual or
after cutting.
alternate bearing, productive, very suscepti-
ble to preharvest drop. Cold-hardy to at least
USES Mainly dessert. Flesh disintegrates
USDA zone 4a (−34°C) (Strang and Stushnoff,
when cooked. Poor to fair for sauce, fair to
1975). ‘McIntosh’ blooms in early-mid sea-
good for juice if mature, usually blended
son, ~5 days before ‘Golden Delicious’. It is
(Smock and Neubert, 1950; Upshall, 1970).
more resistant to bloom frost than ‘Golden
Delicious’, ‘Delicious’ or ‘Jonagold’, but less Low susceptibility to browning after heat
resistant than ‘Cox’s Orange Pippin’ (Sjöstedt treatment for minimally processed slices
(1978), as cited by Nybom, 1992; Mittelstadt, (Kim et al., 1993a).
1989). The lethal temperature resulting in
50% death of flowers was estimated as −3.1°C 4.2.8.3 Storage and postharvest
(Mittelstadt and Salzer, 1989). ‘McIntosh’ has
a high chilling requirement, ~1300 units ‘McIntosh’ stores 2–3 months in air (Smock
(Ghariani and Stebbins, 1994). Tests in the and Neubert, 1950), limited by susceptibility
former USSR suggest that ‘McIntosh’ is fairly to flesh softening, scald and chilling sensi-
resistant to heat and drought (Khalin and tivity. It develops internal disorders (core
Shcherbatko, 1990). flush, brown heart) below 2°C and can also
become mealy in storage (Smock and cultivars (Noiton and Alspach, 1996). It
Neubert, 1950). The range of optimum con- has been used for breeding mainly in
ditions in CA is 1.5–4.5% O2, 1.0–5.0% CO2, Canada, the USA and eastern Europe
1.7–3.0°C, for storage of 5–8 months (including Russia), with some use in
(Kupferman, 1997). Scald control measures the UK. Offspring include ‘Lobo’,
include DPA and air separation/N2 purges ‘Melba’, ‘Summerred’, ‘Spartan’, ‘Cortland’,
in CA. High CO2 injury can occur above ‘Empire’, ‘Macoun’ and ‘Tydeman’s Red’.
10% CO2 (Watada and Abbott, 1985). While some of these have substantial
‘McIntosh’ is susceptible to Coprinus rots in regional importance, none has yet
storage (Meheriuk and McPhee, 1984). surpassed ‘McIntosh’ in commercial trade
volume. The scab-resistant cultivars
‘Liberty’, ‘McShay’, ‘Belmac’, ‘Richelieu’,
‘Enterprise’, ‘Murray’ and many others also
‘McIntosh’ is not prone to sunburn. It has count ‘McIntosh’ in their parentage.
poor colour if night temperatures are too Columnar apples with ‘Wijcik McIntosh’ in
high or nitrogen fertilization is excessive, and their parentage include ‘Telamon’, ‘Tuscan’,
is highly susceptible to preharvest drop. ‘Trajan’, ‘Golden Sentinel’, ‘Scarlett
‘McIntosh’ is highly susceptible to scab; Sentinel’ and others from various European
up to 100% of the fruit may be unmar- breeding programmes. Breeders from the
ketable due to scab if unsprayed (Ellis et al., former USSR reported that ‘McIntosh’
1998). Susceptibility to fire blight and pow- shows high general combining ability for
dery mildew is rated as low to moderate, winter-hardiness (Savel’ev and Yakovlev,
depending on the region. ‘McIntosh’ is 1981).
resistant to cedar-apple rust, quince rust
and hawthorn rust (Aldwinckle, 1974;
4.2.8.6 Sports
Warner, 1992). It is also resistant to race 2
but not race 1 of apple rust in Japan Standard-habit strains have been selected
(Sakuma, 1985). ‘McIntosh’ is susceptible to for improved colour, earlier colour and/or
a number of other fungal diseases, includ- earlier maturity and better storage, e.g.
ing Nectria canker (Bultitude, 1983; ‘Rogers Red McIntosh’, ‘Cornell McIntosh’,
Grabowski, 1994; Braun, 1997), brown rot ‘Summerland McIntosh’, ‘Imperial All Red
(Cimanowski and Pietrzak, 1991) and black- McIntosh’, ‘Marshall Mac’, ‘Redmax®’. They
rot canker (Miller, 1973). However, it shows vary in stability, vigour, yield, quality and
partial resistance to Pezicula bark rots tendency for preharvest drop (Dzieciol et
(Kucmierz et al., 1985) and Alternaria leaf al., 1988; Kruczynska et al., 1991). ‘Pioneer
blotch (Sawamura, 1972) and good resis- Mac’ is not a sport, but an open-pollinated
tance to brown leaf spot (Koropatyuk, seedling of ‘McIntosh’ (Brooks and Olmo,
1974). It expresses only mild symptoms 1997). Several spur types have occurred as
when bud-inoculated with apple mosaic spontaneous limb mutations, e.g. ‘Mor-spur
virus (Singh et al., 1981). McIntosh’, ‘MacSpur McIntosh’, ‘Dewar
‘McIntosh’ is heterozygous for a gene McIntosh’ (‘Starkspur Ultramac’), ‘Wijcik
involved in resistance to rosy leaf-curling McIntosh’ (‘Starkspur Compact Mac’).
aphid (Alston and Briggs, 1977). It has a ‘Wijcik McIntosh’ is unique not only for its
degree of resistance to apple leaf miner columnar shape (classic type 1 in the
(Maciesiak, 1996) and apple rust mite Lespinasse system), but because it is the
(Kozlowski and Boczek, 1987), but is suscepti- only known mutation of apple capable of
ble to red mite (Bielak and Dabrowski, 1986). transmitting compact spurry habit to its off-
spring. The trait is believed to be deter-
mined by a single dominant gene, with
4.2.8.5 Breeding
some modifiers (Lapins, 1974, 1976).
‘McIntosh’ is one of the top five most fre- Commercially, standard types are more
quent founding clones of modern apple popular than the spur mutants.
4.2.9 ‘Rome Beauty’ moderately conspicuous green, brown or

white dots (Beach et al., 1905a; Bultitude,
‘Rome Beauty’, a chance seedling of 1983). Flesh nearly white with green tinge,
unknown parentage, was discovered by H.N. firm, slightly to moderately juicy, a little
Gillett of Proctorville, Ohio, USA, in 1816 coarse, mealy if overripe. Flavour sweet
(Mowry, 1970). ‘Rome Beauty’ was named subacid, slightly aromatic, quality mediocre,
after the township in which it was discovered sometimes lacking in flavour. Exceptional
and was introduced in 1848. Production is ability to withstand handling.
declining in most areas, but it is still impor-
tant in certain regions, such as New York USES See Smock and Neubert (1950) and
State and southern Europe. Synonyms: ‘Gallia Root (1996). Fair for dessert, good for culi-
Beauty’, ‘Belle de Rome’, ‘Faust’s Rome nary use, good for dehydration, fair to good
Beauty’, ‘Gillett’s Seedling’, ‘Morgenduft’. for juice and processing. Sauce of poor
colour and runny, usually blended with
other cultivars (Root, 1996). Good shape and
4.2.9.1 Tree
uniformity for mechanical peeling.
Moderately vigorous, initially upright,
becoming spreading and drooping, terminal
bearing, does not spur readily. Acrotonic tip-
bearing habit, type IV in the Lespinasse sys- Storage and shelf life are short for a late-har-
tem. Much blind wood. Moderately vest apple, 4–5 months in air. ‘Rome Beauty’
precocious, annual bearing, productive, has a strong tendency to become mealy in
diploid, slightly self-fertile. Not subject to storage. Recommended CA conditions are
preharvest drop. Moderately cold-hardy 1% O2, 0% CO2, 0°C for storage of 7–9
(Khanizadeh and Cousineau, 1998; Luby et months (Kupferman, 1997). ‘Rome Beauty’
al., 1999). High chilling requirement, may get superficial scald; it is controlled
~800–1100 h (Tabuenca and Jimenez, 1984). commercially with DPA, low O2, or air
Blooms late season, usually escaping frost. separation/N2 purges of the CA room
(Kupferman, 1997). ‘Jonathan spot’ is elimi-
nated by CA (Smock and Neubert, 1950).
4.2.9.2 Fruit
See Beach et al. (1905a), Mowry (1970),
Bultitude (1983) and Khanizadeh and
Cousineau (1998). Harvested 3 weeks after ‘Rome Beauty’ shows moderate to high sus-
‘Delicious’, 160–175 DAFB (Mowry, 1970; ceptibility to apple scab, fire blight, powdery
Khanizadeh and Cousineau, 1998) in a single mildew and cedar-apple rust. It is moder-
pick. Size usually large; round to round- ately susceptible to quince rust (Aldwinckle,
conic, sometimes slightly oblong or oblate, 1974; Warner, 1992) and susceptible to black-
slightly flattened at base (Plate 4.9). Size and rot canker/frog-eye leaf spot (Miller, 1973). It
shape uniform. Regular or faintly ribbed. is resistant or highly resistant to bitter rot,
Stem cavity smooth and russet-free, wide, Nectria canker (Khanizadeh and Cousineau,
moderately shallow to fairly deep, often gen- 1998) and crown rot (Aldwinckle et al., 1975).
tly furrowed. Stems long, often inserted at an Fruit finish is adversely affected by high pre-
angle, well exserted from fruit base. Calyx cipitation and high temperature (Mowry,
basin medium in width, shallow to moder- 1970). ‘Rome Beauty’ is prone to scarf skin,
ately deep, slight traces of ribs, russet-free. which is particularly noticeable on red
Eye partly to fully open. Skin thick, tough, strains. It is not prone to sunburn. ‘Rome
smooth, glossy, becoming greasy. Ground Beauty’ is more susceptible than ‘Delicious’
colour yellow-green, becoming pale yellow. or ‘Golden Delicious’ to fruit injury from
Original strain 50–80% striped and mottled tufted apple-bud moth (Knight and Hull,
bright orange-scarlet red with some broken, 1988). It has a degree of resistance to woolly
fairly wide carmine stripes. Lenticels small, apple aphid (Asante, 1994).
4.2.9.5 Breeding bearing. ‘Gala’ and ‘Fuji’ may yield more

(Stebbins, 1990). Chilling requirement
‘Rome Beauty’ offspring include ‘Jerseyred’,
medium, ~1140 units (Ghariani and Stebbins,
‘Ben Hur’, ‘Roanoke’ and other cultivars of
1994). Blooms profusely in mid-season;
minor importance (Mowry, 1970). It appears
diploid; not self-fertile but sets heavily.
frequently in the parentages of modern
Blooms on 1-year-old wood and spurs. Very
apple cultivars, although it is not in the top sensitive to chemical thinners (Waliser, 1994).
five founding clones (Noiton and Alspach,
1996). Probably its most lasting breeding
legacy is in scab-resistant cultivars. ‘Rome 4.2.10.2 Fruit
Beauty’ was the first parent crossed with Harvested 150–170 DAFB, about a week
Malus floribunda 821 in the scheme used to before ‘Fuji’ or ‘Rome Beauty’, in several
bring the Vf scab-resistance gene into com- picks (Khanizadeh and Cousineau, 1998).
mercial apple cultivars. All offspring of this Not susceptible to preharvest drop. Size
scab-resistant line therefore carry ‘Rome medium to large; conic to round-conic, more
Beauty’ genes. elongated in cooler regions, with definite
crowning at apex (Plate 4.10). Stem cavity
4.2.9.6 Sports medium in width and depth, usually partly
lined with green or brown russet. Stems
‘Cowin Rome Beauty’, ‘Law Red Rome’, medium to medium-long, even to exserted.
‘Starkspur Red Rome’, ‘Taylor Flamespur’, Calyx basin wide, medium in depth, dis-
‘Cowin Red Rome’ (‘Double Red Rome’) tinctly ribbed. Eye closed or a little open.
(tetraploid). Most strains have improved Lenticels small, conspicuous tan or green
colour; a few are slightly spurrier, e.g. dots. Skin glossy, greenish yellow with short,
‘Lawspur® Rome’, ‘Compact Red Rome’. dark crimson stripes overlaid with dark scar-
Strains vary in fruit shape (Brooks and let blush. Only well-exposed fruit have good
Olmo, 1997), tree size, spurriness, propensity colour. Flesh cream, very juicy, very firm and
to flower on 1-year-old wood, fruit size, crisp, slightly coarse. Flavour sprightly tart-
flower-cluster density, susceptibility to scarf sweet, pleasantly aromatic, refreshing. Flesh
skin (Smith, 1971; Ferree, 1994). browns slowly when cut (Khanizadeh and
Cousineau, 1998).
4.2.10 ‘Braeburn’ USES Appears to be limited to fresh market

at this time.
‘Braeburn’ is a chance seedling introduced by
O. Moran of Nelson, New Zealand, in 1952.
4.2.10.3. Storage and postharvest
Its parentage may be ‘Lady Hamilton’ OP
(Brooks and Olmo, 1997) or ‘Lady Hamilton’ ‘Braeburn’ is susceptible to bitter pit and
× ‘Granny Smith’ (Khanizadeh and other calcium-related disorders in ‘off’ years.
Cousineau, 1998). New Zealand is the world’s It is also prone to water-core, core browning,
main producer of ‘Braeburn’; other producers lenticel blotch and superficial scald. The lat-
are Argentina and Chile. Production is ter is controlled with DPA or ULO storage
increasing in the USA and Europe. (Kupferman, 1997). ‘Braeburn’ will keep for
130 days in air storage (Khanizadeh and
Cousineau, 1998). CA recommendations are
4.2.10.1 Tree
strongly influenced by the prevention of
Low to moderate vigour, very spurry, ‘Braeburn browning disorder’ (BBD), a CO2-
spreading, type II in the Lespinasse system. related severe internal browning. The symp-
Extremely precocious, capable of flowering toms (dry or water-soaked cortex browning
in year of planting and susceptible to stunt- and dry, lens-shaped, internal cavities)
ing if cropped then. Productive, suitable for develop in the first 2 weeks of CA or even on
high-density planting, but can be biennial the tree and are not always visible externally
(Elgar et al., 1998). Texture remains firm. the USA. Useful descriptions appear in Götz
Preharvest causes are not understood. The and Silbereisen (1989), Friedrich and Petzold
incidence rises with high CO2 or low O2 par- (1993), Manhart (1995), Khanizadeh and
tial pressure in CA, rapid imposition of CA, Cousineau (1998) and Anon. (1999). Synonym:
cooler climates/seasons or higher altitude of ‘Lustre Elstar™’.
orchard, advanced maturity, lightly cropped
trees, and other unidentified factors (Elgar et
4.2.11.1 Tree
al., 1998; 1999; Lau, 1998). BBD is not a low-
temperature injury (Lau, 1998) or a mineral- Vigorous, with a strong central leader,
nutrient disorder (Elgar et al., 1999), but may spreading, medium-thick branches, with
be related to the cultivar’s unusually high many water shoots causing a dense, bushy
flesh density (Lau, 1998). Storing ‘Braeburn’ tree; naturally well feathered, intermediate
in air at 0°C for 2 weeks prior to CA appears between type II and III in the Lespinasse sys-
to reduce BBD (Elgar et al., 1998). Optimum tem. Leaves medium to large, very late leaf
conditions vary with region, ranging from 1.0 fall. Precocious, productive, strong tendency
to 3.0% O2, <0.5 to 1.5% CO2, −0.5 to 1.5°C, to biennial bearing. Bears on 2- and 1-year-
for 6–9 months of storage (Kupferman, 1997). old wood and somewhat on spurs. Blooms
mid-season, diploid with fertile pollen,
somewhat self-fertile, but cross-pollination
required. Suitable pollinizers are listed in
‘Braeburn’ is susceptible to scab, powdery Kemp (1996). Medium susceptibility to
mildew and fire blight. It is susceptible to bloom frost. Bark and shoot tips susceptible
apple scar skin viroid (Desvignes et al., 1998). or very susceptible to winter injury
It is very susceptible to red mite. Other dis- (Goddrie, 1985; Fankhauser and Stadler,
ease and insect reactions have not yet been 1986), the latter due to the late cessation of
reported. vegetative growth. Chilling requirement
~1200 units (Ghariani and Stebbins, 1994).
4.2.10.5 Breeding
4.2.11.2 Fruit
‘Braeburn’ is currently in use for breeding in
New Zealand, North America and elsewhere. Harvested 130–150 DAFB, multiple pick.
‘Scifresh’ is an offspring of ‘Braeburn’. Picks fairly easily, very little preharvest drop.
Flesh creamy white to yellow, sometimes
slightly greenish, crisp and fairly firm, juicy,
4.2.10.6 Sports
fine- to medium-textured, balanced sweet-
Recent red sports include ‘Hillwell (Hidala) tart apple with pleasant strong aromatic
Red Braeburn’, ‘Lochbuie™ Braeburn’, flavour. Acidic at harvest, later more mellow.
‘Joburn™ Braeburn’ and others. All offer Usually symmetrical and regular, round-con-
improved red colour and some are claimed ical, sometimes slightly flattened (Plate 4.11).
to ripen earlier. With good light exposure, an attractive,
somewhat orange, bright red, sometimes
slightly pinkish, striped to solid blush on a
4.2.11. ‘Elstar’ green-yellow background, becoming yellow.
Firmness, crispness and taste strongly
Selected by T. Visser and others, Wageningen, depend on proper harvest date. Usually
The Netherlands, from a cross of ‘Ingrid some russet in basin and stem cavity, the lat-
Marie’ × ‘Golden Delicious’ made in 1955, ter often streaking over shoulder. In some
‘Elstar’ was named in 1972 and introduced years, severe netted to solid, rather coarse,
commercially in 1975. It is the main cultivar russet on cheeks (Wagenmakers, 1999). Basin
in The Netherlands and is also planted in medium in depth and width, ribbing fine to
Germany, Belgium, Italy, Denmark, France, medium, apex slightly ribbed. Eye medium,
England, Switzerland, Austria, Poland and half open. Stem cavity rather wide and quite
deep, with medium to long, rather slender 4.2.11.4 Production notes

but sometimes partly thick stalk; fleshy
In areas with small preharvest diurnal tem-
stalks and twinned fruits can occur. Skin
perature differences, poor colour is common.
almost smooth, rather dull, somewhat waxy,
Summer pruning, approximately 2 weeks
moderately greasy with full ripeness,
before harvest, enhances blush. Redder
medium thick, rather tough. Low bruising
susceptibility. Shelf life fairly good. sports can also prevent this problem, but
‘Elstar’ is still a cultivar best suited to cool cli-
USES Especially good for fresh consumption; mates; elsewhere fruit will be too soft. For
good for salads, juice and baking, only mod- good storage and colour, repeated selective
erately suitable for sauce. picking is required. In low-crop years, a leaf
condition resembling Cox’s disease can occur,
in which many shoots have light green
4.2.11.3 Storage and postharvest leaves. Accurate and timely thinning is essen-
‘Elstar’ stores at 1°C for 3 months in air or tial to prevent biennial bearing (Wertheim,
for 5 months at 1°C in scrubbed CA where 1998; Wertheim et al., 2001) and for optimum
both CO2 and O2 are not above 3%. It can be size and taste. Russet can be reduced with
stored in ULO at 1°C for 7–7.5 months at GA4+7. Late leaf fall and soft shoot tips
1.2% O2 and 2–3% CO2, provided picked at increase scab risk; late sprays against Nectria
proper time. Ripe fruit from susceptible canker are required due to late leaf fall. Fruit
plots is stored at 2°C because of the risk of from open trees are less prone to skin spots
internal browning. ‘Elstar’ is susceptible to and fruit from vigorous or poor-yielding
shrivelling, slightly susceptible to bitter pit trees are more prone to internal browning.
and moderately susceptible to scald in high ‘Elstar’ is susceptible to mildew, scab and
CO2. LTB and internal browning (Verschoor Nectria canker (Kemp and van Dieren, 1998).
and de Jager, 1998) can occur, the latter In north-western Europe, mildew suscepti-
enhanced by low storage temperature, cer- bility is no worse than in ‘Jonagold’, but in
tain growing sites, late picking, long storage, the USA the opposite is true. ‘Elstar’ is toler-
high CO2, high relative humidity and plenti- ant of apple scar skin viroid (Desvignes et al.,
ful blush. Red mutants seem to be more 1998) and susceptible to brown rot, crown rot
prone, as do lateral fruits from 1-year-old and sunburn. Some June-drop fruit remain
wood (van der Laken and Verschoor, 1998; on the tree as mummies (Knoche et al., 2000).
van Schaik and Schoorl, 1999). Excessive
internal browning seems to be mainly due to
4.2.11.5 Breeding
storage below 0–0.5°C (van Schaik and
Schoorl, 1999). ‘Elstar’ has been used as a parent in The
In long-term storage, ‘Elstar’ is suscepti- Netherlands (Janse and Verhaegh, 1990) and
ble to schilvlekjes (skin spots), which seem to also by German, French, Italian and Swiss
be related to scald. The occurrence varies breeders. The scab-resistant (Vf ) cultivars
with year. The spots, limited to the surface ‘Ecolette’ (also resistant to powdery mildew),
but spreading over the fruit, occur mainly on ‘Santana’, ‘Gerlinde’, ‘Ahrista’ and ‘Dalinbel’
the shaded side of the fruit. Relatively (‘DL 11’) have resulted, as well as a series of
unripe, poorly coloured fruit from the inner promising numbered selections.
canopy are more prone; growing site also has
an influence. DPA can reduce the incidence
4.2.11.6 Sports
(Roelofs, 1997). A relatively high level of
moisture removal during storage seems to be ‘Elstar’ mutates easily. Since the first sport,
effective for control; high humidity exacer- later called ‘Red Elstar’, was found by Mr L.
bates the problem (van der Valk and Michielsens (Rilland-Bath, The Netherlands,
Tomassen, 1999). Susceptible lots are best 1981), an ongoing series has been tested and
segregated for short storage at low tempera- introduced, mainly for colour pattern and
ture: 0.5°C, 1.2% O2 and 0.5% CO2. intensity, but also growth habit, fruit shape,
firmness and ripening time (Goddrie, 1996). 4.2.12.2 Fruit

Goddrie and Kemp (1992) and Kemp et al.
Harvested 130–150 DAFB, picks easily,
(1994) list over 40 sports and Stehr and
Clever (1995) list 25, but over 75 are now some preharvest drop. Needs full ripeness
known. for optimum taste. Firmness, crispness and
taste strongly depend on proper harvest
date. Size medium, with a slightly brown-
4.2.12 ‘Cox’s Orange Pippin’ ish, orange-red blush and broken stripes
over a pale yellow-green background,
‘Cox’s Orange Pippin’ originated in 1825 on becoming clear yellow. Colour optimum
the property of Richard Cox, in with some cool nights shortly before har-
Buckinghamshire, UK. It is believed to be a vest. Often russeted, varying with year;
seedling of ‘Ribston Pippin’. ‘Cox’s Orange dots and patches of fine to coarse russet,
Pippin’ was introduced in 1850 and was first sometimes with cracks. Sometimes consid-
grown commercially about 1862. In the early erable russet in basin and stem cavity, occa-
1900s, its popularity waned due to disease sus- sionally streaking out. Usually dull, dry
ceptibility, but, after the introduction of lime skin, sometimes medium waxy, fairly
sulphur in the 1920s, it regained commercial smooth. Shape round-conical to oblate, usu-
interest and, since the 1970s, it has been the ally symmetrical (Plate 4.12). Slightly ribbed
most important English apple. It is also grown at basin and apex. Eye fairly small to
in The Netherlands, Germany, Belgium, New medium, half open, calyx closed. Basin
Zealand, Australia, France, Sweden, Denmark medium wide, rather shallow. Stems usu-
and Switzerland. Good descriptions can be ally medium thick, rather short, in a fairly
found in Baldini and Sansavini (1967), Nilsson broad and medium deep cavity; king fruits
(1987), Sanders, (1988), Götz and Silbereisen often have a fleshy stalk. Flesh greenish yel-
(1989), Friedrich and Petzold (1993), Morgan low to deep cream, juicy, firm and crisp,
and Richards (1993), Manhart (1995) and fine-textured; low bruising susceptibility.
Khanizadeh and Cousineau (1998). Synonyms: Sweet, slightly subacid, with a characteris-
‘Cox(‘s) Orange’, ‘Cox’s Orangen Reinette’, tic, rich, aromatic, complex flavour, with
’Kemp’s Orange’, ‘Orange de Cox’ and many notes of nut, pear and banana. Shelf life
others (see Smith, 1971). good. Skin moderately thick, slightly tough.
4.2.12.1 Tree USES Very good for dessert and cider;

preferably blended with others for juice and
Moderately vigorous to vigorous, rather
sauce; unsuitable for salad and pie.
open, naturally very well feathered, spread-
ing-upright, rather slender, long wood. Bears
on 2-year-old wood, spurs and 1-year-old 4.2.12.3 Storage and postharvest
wood. Type III in the Lespinasse system.
Precocious, moderately productive, medium Large fruit from poorly cropped trees are
tendency to biennial bearing; thinning almost unsuitable for storage. ‘Cox’s Orange
required for sufficient size and to prevent Pippin’ stores for 3 months at 3.5°C in air. It
biennial bearing (Child et al., 1986; is susceptible to LTB below 3.5°C and CO2
Wertheim, 1986; Tromp, 2000). Leaves often damage above 1%. ‘Cox’s Orange Pippin’
with necrotic spots; early leaf fall can occur, can be stored in scrubbed CA at 3.5°C, < 1%
especially after a warm, dry period. Blooms CO2 and < 3% O2 for 5 months. Stores in
mid-season, diploid, fertile pollen. ULO 1.2–4% O2, < 0.7% CO2 at 4°C for 6.5
Somewhat self-fertile (Hall and Crane, 1933), months. ULO is preferred to reduce brown
but cross-pollination required. Suitable core. It is susceptible to bitter pit, softening,
pollinizers are listed by Kemp (1996). internal browning, water core, brown core,
Susceptible to bloom frost, very susceptible LTB and shrivelling. Scald incidence varies
to winter injury. Chilling requirement ~1300 from low to moderate, depending on year.
units (Ghariani and Stebbins, 1994). Early picking increases bitter pit and LTB.
4.2.12.4 Production notes (probably) and ‘Kidd’s Orange Red’. It has

also been used in France and Germany.
Best tree condition, cropping and fruit qual-
‘Elstar’ and the scab-resistant American cul-
ity are achieved with good soil and where
tivar ‘Suncrisp®’ are F2 offspring.
summers are cool; it reacts very negatively to
all disturbances in climate, water and nutri-
tion. Especially on weaker rootstocks, Cox’s 4.2.12.6 Sports
disease can occur: mid-shoot necrotic leaf
Götz and Silbereisen (1989) list over 40
spot followed by leaf fall, reddish-purple sports and mutants, and more than 30 are
leaves, blind wood formation and reduced mentioned by Kemp et al. (1998), mainly
productivity. This condition can be reduced based on colour. The English irradiation pro-
by an interstem and stem scoring. Interstems gramme in the 1970s produced many
are also effective against collar rot. Standard mutants, among them some self-fertile
‘Cox’s Orange Pippin’ often lacks colour. ‘Queen Cox’ clones (Campbell and Lacey,
Necrotic leaf spot and leaf fall can be 1982); other irradiated clones came from
reduced by leaf application of magnesium Germany (Götz and Silbereisen, 1989) and
and manganese. Cross-pollination New Zealand (Manhart, 1995). In The
(Goldschmidt-Reischel, 1996) and proper Netherlands, a heat-treated virus-free clone
thinning (Wertheim, 1986) are required for of ‘Cox’s Orange Pippin’ called ‘T12’ (top
adequate size and regular cropping. Severe graft) was introduced in 1975 and is consid-
June drop can occur (Wertheim, 1973). GA ered the standard ‘Cox’s Orange Pippin’.
sprays can induce parthenocarpic fruit Well-known sports include: ‘Cherry Cox’,
(Kotob and Schwabe, 1971). Russeting and ‘Cox Rouge des Flandres’, ‘Crimson Cox’,
rain cracking can be reduced by GA4+7. ‘Queen Cox’, ‘Korallo’, ‘Kortegård Cox’,
Several weekly sprays with calcium are ‘Kummer Cox’, ‘Ottensen Cox’, ‘Moje Cox’,
required to prevent softening and bitter pit ‘Hauschildt Cox’, ‘Ley 36.72’, ‘Clone 18’,
(Sharples and Johnson, 1977). It is suscepti- ‘Cox la Vera’, ‘Red Cox’ and ‘Flikweert’.
ble to damage from sulphur sprays.
‘Cox’s Orange Pippin’ is susceptible to
mildew, collar rot, brown rot and bitter rot, 4.3 Outlook
moderately susceptible to scab (Kemp and
van Dieren, 1998) and very susceptible to World apple production is predicted to rise
Nectria canker, and therefore not suitable for faster than population growth in the near
cold, wet areas. It has low susceptibility to future (O’Rourke, 1998a). Massive produc-
rosy apple aphid and rosy leaf-curling aphid tion in China and new cultivars will have a
(Alston and Briggs, 1977; Graf et al., 1992). major global impact in the coming century.
The key ‘new’ cultivars (‘Fuji’, ‘Gala’,
‘Braeburn’) are expected to increase in pro-
4.2.12.5 Breeding
duction and trade volume over time as
Because of its characteristic aroma, ‘Cox’s more high-density plantings come into full
Orange Pippin’ has often been used in breed- production. ‘Delicious’, ‘Golden Delicious’,
ing and is one of the top five most frequent ‘Cox’s Orange Pippin’, ‘Rome Beauty’,
founding clones of modern apples (Noiton ‘Jonathan’, ‘McIntosh’ and perhaps
and Alspach, 1996). Offspring include ‘Granny Smith’ are losing ground to these
‘Fiesta’ (‘Red Pippin®’) and ‘Meridian’ from and other new cultivars because of changes
the UK, ‘Elise’ (‘Roblos®’), ‘Karmijn de in market demand and consumer prefer-
Sonnaville’, ‘Ivette’ and ‘Zoete Oranje’ (a ence (O’Rourke, 1998b). With the exception
low-acid cooking apple) from The of ‘Braeburn’, these rising cultivars are all
Netherlands, the Czech cultivar ‘Šampion’ the products of breeding programmes.
and the Yugoslavian ‘Pohorka’. Smith (1971) Some groups are moving to control the
lists 110 offspring, including ‘Winston’, availability of their cultivars to limit pro-
‘Holsteiner Cox’ (‘Holstein’), ‘Ingrid Marie’ duction and maintain fruit prices (e.g. in
‘variety clubs’). Disease-resistant cultivars France), ‘Topaz’ (Vf, ‘Rubin’ × ‘Vanda’, Czech
have yet to achieve any major commercial Republic), ‘Pacific Rose’ (‘Gala’ × ‘Splendour’,
importance. Probably the increasing New Zealand), ‘GoldRush’ (Vf, ‘Golden
emphasis on pesticide reduction and Delicious’ × ‘Co-op 17’ USA) and ‘Ambrosia’
heightened efforts of breeding programmes (chance seedling, Canada). Whether they will
to combine good eating quality with resis- ultimately displace major cultivars now in the
tance will change this trend in coming ground remains to be seen.
decades, at least in the European Union.
Cultivars of increasing interest that are
being grown in more than one country now Acknowledgements
include: ‘Cripps Pink’/’Pink Lady®’ (‘Lady
Williams’ × ‘Golden Delicious’, Australia), We are grateful to Richard MacDonald,
‘Honeycrisp™’ (unknown parentage, USA), Reinhold Stainer and Ken Haddrell for read-
‘Corail’/’Pinova™’ (‘Clivia’ × ‘Golden ing the manuscript, to Jean-Marie Lespinasse
Delicious’, Germany), ‘Delblush’/’Tentation®’ for helpful discussions and to Bruce Barritt
(‘Golden Delicious’ × ‘Blushing Golden’, for providing several photographs.
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5 Apple Rootstocks
Anthony D. Webster1 and S.J. Wertheim2

1Crop Science Department, Horticulture Research International, East Malling, West
Malling, Kent, UK; 2Fruit Research Station, Randwijk, The Netherlands
5.1 Introduction 92
5.2 Species, Cultivars and Sub-clones Used as Rootstocks for Apples 92
5.3 Methods of Rootstock Propagation 93
5.4 Rootstock Effects on the Growth and Cropping and the Adaptability to Environmental
Conditions of Scions 94
5.4.1 Rootstocks and interstocks as aids to controlling scion vigour and cropping 94
5.4.2 Rootstocks for adapting scion cultivars to unfavourable environmental conditions 95
5.4.3 Rootstock interactions with scion cultivar and environmental conditions 95
5.5 Rootstock Mechanisms – How do Rootstocks Bring about their Many Effects on Scion Growth
and Cropping? 96
5.5.1 Effects on vigour of growth, yields and fruit size and quality 96
5.5.2 Effects on tree sensitivity to environmental conditions causing severe stress 98
5.6 Breeding New Apple Rootstocks 99
5.7 Choosing the Appropriate Apple Rootstock 100
5.7.1 Attributes of the ideal rootstock 101
5.8 Apple Rootstocks Propagated from Seed 102
5.9 Vegetatively Propagated Rootstock Clones and Sub-clones Used to Control Tree Vigour 102
5.9.1 Super dwarfing selections 103
5.9.2 Dwarfing selections 103
5.9.3 Semi-dwarfing selections 107
5.9.4 Semi-vigorous to vigorous selections 111
5.9.5 Very vigorous selections 111
5.10 Clonal Rootstocks Used to Adapt Trees to Unfavourable Environmental Conditions 111
5.10.1 Tolerance to winter-cold injury 111
5.10.2 Tolerance to soil-borne or aerial pathogens 118
5.10.3 Tolerance to soil-borne or aerial pests 118
5.10.4 Tolerance to drought or soil asphyxiation 119
5.11 Use of Interstocks and Interstems 119

92 A.D. Webster and S.J. Wertheim
5.1 Introduction unsuccessful when used for propagating

apple-scion cultivars. In contrast, the root-
A rootstock constitutes the root system and a stocks used for apple tree propagation can in
small proportion of the lower trunk (some- most instances be propagated easily and
times referred to as the shank) of most apple cheaply, either from seed or from simple lay-
trees. Grafting the genetically distinct fruiting ering or cutting techniques. These rootstocks
part of the tree, the scion, on to the rootstock are then used to multiply selected scion cul-
forms the whole tree (or stion). Occasionally, tivars of apples by employing grafting or
a third genetically distinct component, an budding techniques (see Chapter 6).
interstock or interstem, is grafted between Although new improved methods of propa-
the rootstock and the scion (Fig. 5.1). gation based on cutting techniques (includ-
It is thought that rootstocks have been ing misting, fogging, heated bins and
used as an aid in the propagation of apple micropropagation) have in recent years led
trees for more than 2000 years. The principal to successful techniques for propagating
reason for their use is the difficulty in propa- apple scions on their own roots (see Chapter
gating selected cultivars of apple scions on 6), these have yet to be adopted by commer-
their own roots. Although apple trees can be cial nurserymen. Rootstocks continue to be
propagated quite easily from seed (pips), the used as a principal aid to the propagation of
resulting trees are extremely variable in apple scions.
vigour, habit and fruit characteristics; most Rootstocks may also confer many other
apple trees raised from seed are vigorous and benefits, in terms of the growth and crop-
bear fruits of poor size, appearance and qual- ping of the scions grafted on them. Selected
ity. This is because of the heterozygous nature rootstocks, especially clonal selections, may
of Malus pumila Mill. Unfortunately, therefore, be used to control the intrinsic vigour of the
trees propagated from seeds collected from an scion, its habit, its precocity and efficiency of
apple tree exhibiting desirable fruiting charac- cropping and the quality of fruits produced.
teristics will not be true to type. Rootstocks may also be used to adapt scions
Although apple scions can with some dif- to unfavourable environmental (climatic and
ficulty be propagated from layers, the tech- soil) conditions and against aerial and soil-
nique is difficult and the multiplication of borne pests and diseases.
scion trees using this technique is extremely Interstocks or interstems are occasionally
slow. Traditional methods of propagation used in propagating apple trees. These are
using cutting techniques are generally usually clonal and genetically distinct from
the scion and the rootstock. They are used
for aiding the control of tree vigour, stimulat-
ing precocious fruiting and improving the
Dwarfing effects
branching of young nursery trees and to
Fruiting scion cv. increase resistance against trunk diseases or
winter cold injury.
Fruiting scion cv.
5.2 Species, Cultivars and Sub-clones Used

as Rootstocks for Apples
Dwarfing
Graft Graft interstock
union Dwarfing union
rootstock Invigorating The apple rootstocks used throughout the
rootstock
world are, like the fruiting apple scions, clas-
sified by taxonomists as M. pumila Mill. (or
Increasing height of
budding/grafting
Use of dwarfing rootstock
clones as interstocks
alternatively as Malus × domestica Borkh.).
increases dwarfing can also dwarf scions. Most plant taxonomists believe that the culti-
of scion The effect is proportional
to the interstock length vated apple and its rootstocks are derived
Fig. 5.1. Influence of height of budding or dwarfing from a hybrid of several species formed cen-
interstock on the vigour of scion growth. turies ago close to Almata (Alma Ata) in
Apple Rootstocks 93
Kazakhstan, where natural forests of Malus as rootstocks for apples (Schmidt, 1988).
predominate. These hybrids were then However, absence of any vigour control and
spread, possibly by animals such as bears difficulties in separating the apomictic from
but almost certainly by humans, along the the non-apomictic seedlings have made this
Silk Route into Persia, where horticulturists rootstock option unpopular with commercial
domesticated them. Later, the superior nurserymen and fruit growers.
and/or useful selections were spread much Horticulturists have, over the last 150
further within the extensive Roman Empire. years or more, bred and selected many culti-
Currently, however, there is little DNA evi- vars of clonal rootstocks. Some of the first
dence to support the contention that the rootstock cultivars were selected as chance
domesticated apple is a hybrid, and some seedlings from wild populations of apples.
botanists now believe that the cultivated More recently, fruit breeders have made con-
apples and their rootstocks are in fact all trolled crosses and selected from the result-
similar to the wild apples, now referred to by ing siblings. All of these rootstock cultivars,
some apple taxonomists as Malus sieversii with the exception of the apomicts men-
(Ledeb.) Roem. tioned above, must be propagated vegeta-
In contrast, the rootstocks used for propa- tively. Selections within the cultivar have
gating many stone fruit species (peaches, apri- also been made, especially with the rootstock
cots, cherries and plums) are of different species M.9, where ‘sub-clones’ differing slightly in
from the fruiting scion species, and some may their individual characteristics have been
be hybrids developed by plant breeders. Even distinguished. Although genetically similar,
a different genus, the quince (Cydonia oblonga these ‘sub-clones’ differ slightly in their abil-
L.), is widely used as a rootstock for European ity to propagate and in their effects upon
pears (Pyrus communis L.). The only species scion vigour.
other than M. pumila (including M. sieversii and
M. × domestica) that is widely used as a root-
stock for apples is Malus prunifolia (Willd.) 5.3 Methods of Rootstock Propagation
Borkh. Seedlings of this species are used as
rootstocks in parts of China. Traditionally, one method of propagating
The use of different Malus species as root- rootstocks was to dig up suckers from
stocks for apples was first suggested in the around the bases of mature apple trees. This
west by Sax (1949), but they had already method is no longer used and is not to be
been used for centuries in eastern countries, recommended due to the possibility of virus
such as China. One of the problems with transmission via the rootstock to the new
using seedling-raised rootstocks has always scion tree. More often, seeds of apples col-
been their rather variable effects on scion lected in the wild or from orchard trees were
growth and cropping. Luckwill and used for propagating rootstocks. Such root-
Campbell (1954) suggested using apomictic stocks, although cheap to propagate, were
seedlings of Malus species to give more uni- mostly very invigorating and also very vari-
formity of performance. Apomictic seeds are able in their effects on scion growth and
derived only from the maternal tissues and cropping. Apple seedlings are still used as
are homozygous and very uniform in perfor- rootstocks in some parts of the world,
mance as rootstocks. Campbell and Wilson although today seedlings of predominantly
(1962) continued this work on apomicts, one cultivar (e.g. ‘Red Delicious’ in the USA)
focusing on Malus hupehensis (Pamp.) Rehd. provide slightly improved uniformity when
and Malus toringoides (Rehd.) Hughes. these rootstocks are grafted with scions.
Apomictic seedlings of hybrids between M. They are of most value where control of the
× domestica and M. hupehensis, Malus sargentii scion vigour and cropping is achieved by
Rehd. or Malus sieboldii (Reg.) Rehd., all of other methods than by using a dwarfing
which are cheap to propagate, are free from rootstock, but they may also be used on
virus and have uniform effects on scion droughty soils where their deep root systems
growth and cropping, have also been tested are of value.
Where rootstocks are needed to provide Carlson, 1987; Wertheim, 1998). Many
additional benefits, other than simply a apple rootstocks/interstocks reduce the
means of propagating the scion, rootstocks strong vigour of scions, allowing them to
propagated using vegetative techniques (i.e. be grown as dwarf, closely planted trees
clonal rootstocks) are essential. These root- that are easy and inexpensive to manage.
stock cultivars, which can affect the vigour, Trees grown on most of these dwarfing
habit and cropping performance of scions as rootstocks/interstocks also produce fruits
well as adapting the scions to unfavourable precociously and crop abundantly and con-
environmental (edaphic and climatic) condi- sistently from season to season (Webster,
tions, are now used for tree propagation in 1994). Studies conducted in several apple-
the majority of apple-producing countries of producing countries of the world have
the world. Their propagation, by stooling, shown that, where the unit costs of labour
layering or cutting techniques (including in and/or land are relatively high, controlling
vitro micropropagation), is described in tree vigour with dwarfing rootstocks/inter-
Chapter 6 and also in Webster (1995). stocks greatly improves the economics of
apple production. Trees of dwarf stature
can also be targeted with crop-protection
5.4 Rootstock Effects on the Growth and sprays accurately, so avoiding excessive use
Cropping and the Adaptability to of pesticides and undesirable spray drift
Environmental Conditions of Scions into the surrounding environment.
Use of appropriate dwarfing root-
Rootstocks capable of influencing scion stocks/interstocks often increases the num-
vigour have been available to horticulturists bers of floral clusters (spur, terminal and
for two millennia at least, there being evi- axillary) produced per linear branch length
dence from ancient Persia of dwarfed apple by the scion (Ferree et al., 1995). The quality
trees grown on rootstocks. However, it is of the flowers produced by these floral
only in the last 100 years that the full poten- buds (i.e. their ability to set fruits when
tial of rootstocks for modifying scion growth pollinated) can also be improved by use of
and cropping and adapting trees to certain rootstocks, although the evidence
unfavourable environmental conditions has for this is often inconsistent from site to
been recognized. This rootstock potential is site. Rootstocks such as M.9 have been
still being developed in many parts of the reported to improve while others, such as
world. For many centuries the majority of M.27, have been reported to reduce the size
apples were produced on large standard of scion fruits at harvest. However, these
trees grown either on their own roots or on effects are also often difficult to measure
rootstocks raised from seed or from suckers objectively, unless crop loading on trees on
dug up from beneath mature orchard trees. the different rootstocks is uniform. It is
Only in the gardens of European monaster- also suggested that rootstocks can influ-
ies, palaces and castles and a few private ence the postharvest storage potential of
gardens were smaller trees on more dwarf- apples, although this effect is also difficult
ing rootstocks produced. to prove conclusively on account of sea-
sonal influences and the confounding
effects of the rootstock on crop load/tree
5.4.1 Rootstocks and interstocks as aids to and time of fruit ripening (Autio, 1991;
controlling scion vigour and cropping Barden and Marini, 1992). It is generally
agreed that scions on M.9 rootstocks or
Choice of the appropriate rootstock and/or interstocks generally ripen up to 1 week
interstock can enable the fruit grower to earlier than the same scion on other more
control the inherent vigour of the scion invigorating rootstocks (Hewetson, 1944),
tree, so making possible and facilitating the and trees on M.27 have also been reported
adoption of a chosen system of tree spac- to ripen earlier than trees on M.26 (Lord
ing, pruning and training (Ferree and et al., 1985).
Apple Rootstocks 95
5.4.2 Rootstocks for adapting scion cultivars be a strong need for a greater selection of
to unfavourable environmental conditions rootstocks that are both drought-tolerant and
dwarfing.
In many areas of the world where apples are Rootstocks also adapt scions to soils
produced commercially the environmental infested with damaging soil pests and/or
conditions are not fully suitable. Very severe pathogens. The woolly apple aphid (Eriosoma
winter cold can lead to the death of apple lanigerum Hausmann) limits apple produc-
trees on many rootstocks and the require- tion significantly in many parts of the world,
ment for rootstocks in many eastern especially in the southern hemisphere. This
European countries, Canada and the USA is pest damages apple roots most severely and
for resistance to low-temperature damage. In in some cases can lead to tree death.
recent years many cold-tolerant selections Rootstock selections, such as the
have become available. Although these resis- Malling–Merton (MM) series are resistant to
tant rootstocks are believed to slightly aid this pest and enable production in areas nor-
scion tolerance to winter cold, their main mally blighted by this pest.
benefits are in providing the roots and Damaging pathogens, such as collar or
trunks (shanks) of the rootstock with greater crown rot (Phytophthora cactorum (Leb. and
tolerance to very low temperatures. Where Cahn) Schroet), also limit apple production
winter temperatures are very low for sus- severely, especially on heavy clay or other
tained periods and snow cover is minimal, poorly draining soils. The popular semi-vig-
rootstocks such as M.9 and M.7 are often orous rootstock MM.106 is particularly sensi-
severely damaged or even killed. In contrast, tive to this problem. In recent years, many
the roots of other rootstocks show better tol- apple rootstocks have been selected that show
erance of freezing soil conditions (Czynczyk, strong tolerance/resistance to this disease.
1974; Quamme, 1990; Quamme et al., 1999). Care must be taken in selecting rootstocks
Sensitivity to drought is a major con- so as not to increase the sensitivity of trees to
straint on apple production in some coun- other damaging pests and diseases. Use of
tries and this sensitivity may be influenced the dwarfing and semi-dwarfing rootstocks
by choice of rootstock (Olien and Lakso, M.9 or M.26 is not to be recommended in
1984; Higgs and Jones, 1991). Sensitivity is areas where the bacterial disease fire blight
particularly severe on dwarfing and very (Erwinia amylovora (Burr.) Winslow et al.) is a
dwarfing rootstocks, such as Mark, P.22 and problem. Both of these rootstocks are partic-
M.27. Some semi-dwarfing rootstocks, such ularly sensitive to this pathogen. In apple-
as J.9, appear to offer some tolerance of production areas where various species of
drought conditions. By selecting rootstocks mice or voles cause severe damage to the
that are more tolerant of drought, apple pro- tree’s root system, rootstocks such as Novole,
duction has been rendered more feasible in which show some resistance to these pests,
many droughty areas. Sometimes the prob- have been considered.
lem is overcome by using very drought-
tolerant but invigorating seedling rootstocks,
as in China. Control of tree vigour and crop- 5.4.3 Rootstock interactions with scion
ping is then achieved by use of a clonal inter- cultivar and environmental conditions
stock, such as M.26. The extensive and deep
root systems of these invigorating seedling The literature concerning the relative effects of
rootstocks could explain their drought toler- different apple rootstocks on scion growth and
ance, although they may also be able to cropping is sometimes inconsistent, leading to
extract tightly bound water from clay soils confusion when seeking the appropriate
better than other more drought-sensitive choice of rootstock. This is best explained by
rootstocks. With the current trend towards the usually subtle, but occasionally significant,
reduced use of herbicides under organic sys- interactions between the rootstocks and the
tems of production and the need to econo- scions (Tubbs, 1980) or the rootstocks and the
mize on water use in many areas, there may environmental conditions (Olien et al., 1991).
Usually, the relative effects of different affect the abundance of flowering, fruit set
rootstocks on the growth and cropping of the and yields, and the mechanisms by which
scion remain similar for the majority of scion these influences are brought about are also
cultivars (Hirst and Ferree, 1995). not understood. Many theories have been
Nevertheless, the inherent vigour of the advanced in attempts to explain the dwarfing
scion cultivar will greatly influence final tree effect on apple scions of some rootstocks
size on any particular rootstock. This is why (Tubbs, 1972; Lockhard and Schneider, 1981).
growers are advised to choose the more Some suggestions, now less fashionable,
dwarfing sub-clones of M.9 or even M.27 sought to explain the dwarfing effect in terms
rootstock for a vigorous scion cultivar such of changes in tree–water relations or nutrient
as ‘Elstar’, while with the less vigorous uptake caused by the rootstock or its imper-
‘Cox’s Orange Pippin’ the more invigorating fect graft union with the scion (Jones, 1971,
sub-clones are recommended. 1975). More recently, scientists have focused
The rootstock’s interaction with environ- on studying changes in the production and
mental (soil and climatic) conditions are often movement of plant hormones within the tree
very significant and frequently the wrong brought about by use of the rootstock
choice of rootstock or tree spacings is made. (Soumelidou et al., 1994; Kamboj et al., 1997).
Until rootstock evaluation trials pay more The hypothesis is that the rootstock, or possi-
attention to the prevailing environmental bly its graft union with the scion, alters the
conditions and endeavour to understand and ratios and concentrations of the growth-pro-
explain these interactions, this inconsistency moting hormones, such as auxins, gib-
of response is likely to remain an enigma. berellins or cytokinins, and maybe also the
When choosing a new rootstock it is always inhibiting hormones, such as abscisic acid,
preferable if small test plantings can be estab- which are translocated within the tree. The
lished on the site prior to any large-scale work by Soumelidou et al. (1994) and Kamboj
commitment to the new rootstock. Where this et al. (1997) suggested that the rates of
is not possible, it is vital to take account of basipetal auxin translocation were less in
the results of trials of the new rootstock that dwarfing than in invigorating rootstock
have been conducted in similar environmen- stems. The later work also indicated that the
tal conditions and with the same scion culti- ratios of abscisic acid to auxin content were
var. Recent studies in the USA have higher in the bark of dwarfing rootstocks and
attempted to use data collected from multi- that differences in cytokinin translocation
site trials to estimate a ‘site index’ to be used rates may also be measurable (Fig. 5.2).
in the construction of predictive models to It is important to note that the dwarfing
aid rootstock selection (Olien et al., 1995). To influence of rootstocks on apple scions is dif-
help apple growers improve their rootstock ferent from that achieved when using most
choices in the future, more research and compact scion cultivars or chemical plant-
information are needed on predicting the growth regulators. In comparison with invig-
growth potential of sites for apple orchards. orating rootstocks, dwarfing rootstocks/
interstocks reduce the speed of extension
shoot growth throughout the season and
5.5 Rootstock Mechanisms – How do often bring about an earlier termination of
Rootstocks Bring about their Many this shoot extension in the late summer or
Effects on Scion Growth and Cropping? early autumn. This effect and changes in tree
habit towards more horizontal branch orien-
5.5.1 Effects on vigour of growth, yields and tation (Warner, 1991) together account for the
fruit size and quality effects of the dwarfing rootstocks in reducing
the size of apple-scion trees. In comparison,
There is still little or no understanding of most compact types of scions and chemical
how rootstocks bring about their effects on plant-growth regulators reduce tree size by
the vigour of shoot growth of scions grafted shortening the internodes of extension
upon them. Choice of rootstock can also shoots. The full implications of these differ-
Apple Rootstocks 97
How do dwarfing rootstocks and/or 50 (a)

root manipulation techniques
bring about their effects on
scion growth and cropping? Mac 9
40 MM.106
M.9
P.16
Length (cm)
30 P.2
Hypothesis
Dwarfing rootstocks
affect production Bud 9
and basipetal 20
translocation M.27
of auxins to roots
Scion cv. 10
0
17/5 31/5 14/6 28/6 12/7 26/7 9/8
24/5 7/6 22/6 5/7 19/7 2/8
80 (b) MM.106
Hypothesis
The discontinuity of xylem
and phloem at the graft
Graft union union changes the fluxes
Hypothesis of hormones, assimilates, 60
The stems (shanks) of water and/or nutrients Mac 9
Length (cm)
dwarfing rootstocks produce between scion and

rootstock M.9
inhibitors or bind growth
promotors moving through Rootstock 40 P.16
them P.2
Bud 9
Hypothesis Hypothesis
Reducing root volume Roots of dwarfing M.27
by pruning or restriction rootstocks produce 20
reduces or increases and export different
hormones, water or amounts of hormones,
nutrient supply to scion water and nutrients
Fig. 5.2. Some hypotheses concerning how apple 0
17/5 31/5 14/6 28/6 12/7 26/7 9/8
rootstocks bring about their dwarfing influence 24/5 7/6 22/6 5/7 19/7 2/8
upon scion vigour. Day/month
Fig. 5.3. Effect of different rootstocks on the growth
ences in mode of dwarfing action on tree of scion extension shoots. (a) Shoots with horizontal
performance and productivity have yet to be orientation. (b) Shoots with vertical orientation.
fully explored. However, short internodes
and much mutual shading of leaves might be comparison with seedling rootstocks inducing
expected to have some negative effects on the same level of scion vigour. Also, use of
cropping and fruit quality in areas with less some interstems, which have no significant
than ideal light interception. This may be influence on tree vigour and size, may also
one contributory reason why compact spur have very beneficial effects on fruiting precoc-
types of cultivars such as ‘Granny Smith’ ity. This is possibly explained by the high
(e.g. ‘Granspur’) often crop less efficiently quality (size and feathering) of trees raised
than the standard non-compact parent and using an interstem rather than the interstem
others, such as ‘McIntosh Wijcik’, are itself (Wertheim and Callesen, 2001).
severely biennial (Fig. 5.3). Reducing root growth (by root pruning or
The beneficial effects of dwarfing root- root restriction) brings about dwarfing
stocks/interstocks on the precocity and effi- effects on scions grown on invigorating root-
ciency of tree yields have often been attributed stocks (Schupp and Ferree, 1987; Webster et
to a change in the partitioning of the dwarfed al., 2000) that are similar to those brought
tree’s assimilates from shoot growth to fruit about by use of dwarfing rootstocks. It could
production. However, this explanation is too be argued, therefore, that the growth reduc-
simplistic, as there are several invigorating tion caused by dwarfing rootstocks might be
and semi-invigorating clonal rootstocks, such partly due to them bringing about a reduc-
as M.25 and MM.106, which also induce tion in the root : shoot ratio of the tree. Most
improved yield precocity and efficiency in dwarfing clones certainly produce smaller
root systems than more invigorating root- growth. This earlier termination in growth
stock clones. However, this hypothesis fails should also result in a change in the biosyn-
to explain the growth reduction caused by thesis and translocation of hormones from
dwarfing rootstock clones when they are the shoot tip. The production of gibberellins
used as interstocks, where the dwarfing and auxins in the meristems and new
clone itself forms no roots. The dwarfing unfolding leaves of actively growing shoots
effect of using a rootstock such as M.9 or and their translocation basipetally undoubt-
M.27 as an interstock increases with the edly diminish once a resting bud is formed
length of interstock used up to approxi- at the apex. Although a partially plausible
mately 25 cm, indicating the influence of hypothesis, it fails to explain why certain
some stem-derived dwarfing factor (Parry invigorating rootstocks, such as M.25, stimu-
and Rogers, 1968). The same phenomenon is late improved flowering and cropping in
noted when scions are budded at different scions but with no reduction in tree vigour
heights on the shanks of dwarfing apple compared with trees on more poorly crop-
rootstocks (Fig. 5.4); trees budded high are ping but equally vigorous rootstocks.
much more dwarfed than those budded Whether M.25, like dwarfing rootstocks, also
close to ground level (Parry, 1976). induces termination of shoot growth early in
One possible explanation is that growth- the season is not known. This is another area
inhibiting substances are formed or growth- of research in need of further study.
promoting substances are broken down
within the stems of dwarfing rootstock
clones and that this either causes a direct 5.5.2 Effects on tree sensitivity to
effect on scion extension growth or alterna- environmental conditions causing severe
tively indirectly influences scion shoot stress
growth by effects on root growth. Much fur-
ther research will be needed before the Little is understood of the mechanisms by
mechanisms by which dwarfing rootstocks which rootstocks differ in their resistance/
operate are understood. tolerance to damaging pests or diseases,
Rootstocks also influence apple-yield pro- severe cold, drought or other unfavourable
ductivity (Rom et al., 1990) but how they do conditions. A few studies conducted recently
this is also poorly understood. One hypothe- have sought to explain the reasons for differ-
sis is that trees on dwarfing rootstocks termi- ences in apple-rootstock sensitivity to
nate shoot growth earlier in the summer drought (e.g. Atkinson et al., 1999). However,
than trees on more invigorating rootstocks most of the rootstock effects on these para-
and thereafter partition more of their avail- meters remain an enigma and more studies
able assimilates towards the sites of floral are needed in this area to aid rootstock
primordia and less towards further shoot breeding and selection in future years.
Scion
Graft union
Scion
Graft union
Rootstock Rootstock
Fig. 5.4. Influence of height of budding on scion vigour.

Apple Rootstocks 99
5.6 Breeding New Apple Rootstocks stock and induced poor yield precocity and
efficiency. In breeding programmes under-
Approximately 100 years ago, all the clonal taken jointly by the John Innes and East
apple rootstocks used by nurserymen for Malling Institutes in the UK, resistant root-
raising apple trees had been selected from stocks, known as the Merton Immune (e.g.
seedling populations and their precise origin Merton 793) and the MM series, were devel-
is, for the most part, uncertain. These clones oped (Tydeman, 1953). Preston (1955, 1966)
had become very mixed up by the turn of the described some of the initial trials using the
last century and apple growers could never MM rootstocks. One promising invigorating
be certain of consistent performance when rootstock, M.25, that did not show resistance
using them. A now well-documented pro- to woolly apple aphid was also released
gramme of research undertaken by East from this breeding programme.
Malling Research Station in England set The popular rootstocks from the original
about sorting out these mixtures and, by selections (M.7 and M.9) together with the
1920, a series of rootstocks had been identi- subsequent releases of M.26, M.27 and
fied, described and distributed to nurseries Merton 793 and the MM series, provided
(Hatton, 1917). Initially, nine types were fruit growers in many parts of the world
released and among these type IX (Jaune de with reliable rootstocks offering a range of
Metz or M.9) and M.7 remain popular scion vigour control. However, these root-
throughout the world today. Further selec- stocks are not suited to all areas of apple pro-
tions followed, but most of these proved duction. M.9 and M.27 are sensitive and very
very invigorating and most have now disap- sensitive to winter cold injury, respectively,
peared from commerce. Descriptions of the and fruit-breeding programmes have been
Malling apple-rootstock selection numbers 1 initiated in many parts of the world to pro-
to 16 are given by Pearl (1932) and details of duce dwarfing rootstocks exhibiting
their effects on tree growth and cropping by improved cold tolerance. This focus on cold
Hatton (1935) and Tydeman (1955). tolerance in apple-rootstock breeding has
Following the early selection work under- been and remains of particular importance in
taken by East Malling, there were only two the programmes conducted in eastern and
rootstocks, M.9 and M.8, that could be central Europe. Details of breeding pro-
classed as dwarfing. The second of these programmes in Poland, Belarus, the Ukraine,
duced very brittle roots and trees on it were Russia, the Czech Republic and the Baltic
very poorly anchored. In attempts to extend states are given in Sadowski and Hrotko
the range of dwarfing rootstocks and to pro- (1999). Breeding for cold tolerance has also
duce more invigorating rootstocks that been a prime objective of programmes in
exhibited good yield precocity, crosses were Canada (Quamme, 1990; Elfving et al., 1993;
made using M.9 as one parent (Tydeman, Quamme et al., 1999) and the USA (Cummins
1933, 1943). The first results of orchard trials, and Aldwinckle, 1995). Usually cold-tolerant
comparing the best of these selections, were cultivars, such as ‘Antonovka’, or hardy crab
reported by Preston (1954). One extremely apples or the cold resistant rootstock A2
dwarfing selection, 3426, was never released have been used in crosses with M.9 or M.8.
but two other selections that showed great Other problems became evident with the
promise, 3436 and 3431, were eventually existing range of rootstocks. Many, including
released as the semi-dwarfing M.26 and the M.9 and M.26, are very sensitive to fire blight
very dwarfing M.27 rootstocks. attacks, while MM.106 is very susceptible to
The woolly apple aphid (E. lanigerum) has collar (crown) rot and both MM.106 and M.26
always caused big problems to apple pro- are sensitive to tomato ringspot virus. Also,
ducers in the southern hemisphere, where it despite the success of the MM series of root-
severely damages the root systems of trees. stocks against woolly apple aphid, none of
Traditionally, the resistant scion cultivar this series is dwarfing. In a comprehensive
‘Northern Spy’ was used as a rootstock. programme of breeding, scientists at Cornell
However, this was a very invigorating root- University in the USA have sought to over-
come these and other problems and produce the correct choice is made to suit the environ-
new improved clones of rootstocks for apples mental conditions, the economic constraints
(Cummins and Aldwinckle, 1995; Johnson, and the management strategies of the partic-
1999; Robinson et al., 1999). Although show- ular apple-production enterprise. Choice of
ing initial promise, most of these Geneva, the appropriate rootstock can, in certain cir-
New York, selections have yet to be fully cumstances, make the difference between
tested in different parts of the world. profitability and loss of the apple orchard.
It is important that scientists and others Unfortunately, rootstock decisions made by
involved in breeding rootstocks take note of growers are not always as objective and ratio-
the changing requirements of apple produc- nal as they might be. The current fashion for
ers throughout the world. As chemical soil the use of a particular rootstock in another
fumigants are withdrawn from use and as area of the world, poor or biased advice from
organic systems of production (under which extension specialists or simply pressure to
weed competition may increase) become sell by nurseries may all lead to the wrong
more popular, there will be a growing need choice being made. In choosing the appropri-
for dwarfing rootstocks that are more able to ate rootstock it is essential to list and rank the
tolerate replant problems and drought stress. priorities needed, based on the environmen-
All new rootstocks have, until recently, tal and other constraints relevant to the fruit
been produced using conventional techniques enterprise and the chosen site for the orchard.
of breeding. In most cases existing rootstocks, The current fashion in many parts of the
such as M.9, have been crossed with other world is to grow apple trees on dwarfing
rootstocks, scion cultivars or Malus species rootstocks, such as M.9. The trees are dwarf
and the siblings screened for the desired char- in stature and hence easier and cheaper to
acteristics. In the future, novel techniques of manage and harvest than larger trees. Trees
fruit breeding, which use new methods of on M.9 crop precociously and abundantly
molecular biology, may play a greater role in and produce fruits of large size. The dwarfed
producing new rootstocks. Already, genes trees also have environmental advantages in
derived from the cecroptin moth have been that they can be targeted with agrochemical
introduced into M.26 and M.9 rootstocks, in sprays very accurately, so greatly reducing
attempts to impart fire blight resistance into spray drift into the surrounding atmosphere.
these susceptible clones (H. Aldwinckle, Considering these advantages, it is easy to
Geneva, New York State, 2000, personal com- appreciate why M.9 has become so popular
munication). Similarly, rol genes have been throughout the world and why many grow-
used in other work in attempts to improve ers might consider no other rootstock when
rooting and dwarfing characteristics of root- planning their new orchard. Nevertheless,
stocks (Welander, 1998; Welander and Zhu, like all other rootstocks, M.9 has several criti-
2000). This research is still at an early stage cal disadvantages. First, it is very sensitive to
and it is not yet known how consistent the damage to its root system by the woolly
expression of the introduced characteristics apple aphid (E. lanigerum), which has, in the
will be once trees are raised on the transgenic past, proved a constraint on production in
rootstock clones and planted in the orchard. many southern hemisphere countries. It is
In Europe, it is also possible that fruits raised also very sensitive to fire blight and orchards
on trees propagated on transgenic rootstocks grafted on to it in the eastern states of the
may prove unacceptable to consumers. USA have recently suffered huge tree losses
as a result of this disease. In addition, M.9,
like most other dwarfing rootstocks, is sensi-
5.7 Choosing the Appropriate Apple tive to winter cold injury, particularly on
Rootstock poorly drained soils, making its use in east-
ern and central Europe, parts of Canada and
A very large range of rootstocks, either the USA very risky. It is also poorly
seedling or clonal, is now available for use by anchored, needing expensive stakes or other
nurserymen and fruit growers. It is vital that means of support in all situations, and it also
Apple Rootstocks 101
tolerates hot, dry soils very poorly. Where healthy. More information on the diseases
land and labour costs are relatively inexpen- affecting apple rootstocks and scions is
sive but the costs of nursery trees are high, it contained in Chapter 18.
may be advantageous, economically, to
The other desirable rootstock attributes
choose a more invigorating rootstock than
are shown below. How these are ranked in
M.9. A similar choice may be appropriate on
order of priority will depend largely on the
soils of lower-than-average fertility or where
specific needs of the nurseryman or fruit
water supplies are very limited on sites sub-
grower and the particular constraints affect-
ject to transient drought. On sites such as
ing his or her tree or fruit production.
these, M.9 and most other dwarfing root-
stocks are inappropriate choices.
The above brief points are made as an 5.7.1.1 Attributes important to the
example to emphasize the importance of nurseryman
careful ranking of priorities when choosing
• Ease of propagation. Rootstocks that are
an apple rootstock.
difficult to propagate are unpopular with
nurserymen and result in trees that are
very expensive to produce. All rootstocks
5.7.1 Attributes of the ideal rootstock
should be easy to propagate from seed or
from layering or cutting techniques.
There are certain attributes common to all
• Good performance in the nursery. The
good clonal (vegetatively propagated) root-
ideal rootstock should establish well in
stocks. The essential attributes for all root-
the liner nursery, exhibit good bud or
stocks are:
graft compatibility with the scion and
• Long-term graft compatibility with the produce well-feathered trees.
scion. No grower can tolerate delayed
incompatibility occurring in the orchard
5.7.1.2 Attributes important to the fruit
and premature tree death. Fortunately,
producer
graft incompatibility is only occasionally
a problem with apples, usually when • Ability to control scion vigour to the
trees are grafted on to species or hybrids required level. The chosen rootstock
of crab apples. should be capable of controlling the
• Good health. Rootstocks used for raising vigour of the scion trees to the level
apple trees should be free from damaging required by the grower. This will be influ-
pests and diseases. Of particular impor- enced by the environmental conditions in
tance is freedom from virus and bacterial the orchard and by the management sys-
diseases. Virus-infected rootstocks will tem adopted by the grower.
transmit the disease to any scions budded • Ability to induce precocious and abundant
or grafted on to them and usually this cropping. The ideal rootstock should
will result in reduced growth, yield and induce scions to flower and crop signifi-
fruit quality. It is vitally important that cantly in the first few years following
rootstocks guaranteed free from viruses planting, if rapid returns on orchard invest-
are chosen. The bacterial disease crown ments are to be achieved. The rootstock
gall (Agrobacterium tumefaciens (Smith and should also induce consistent and abun-
Townsend) Conn.) reduces scion-tree dant cropping of large high-quality fruits.
growth on some soil types and most • Resistance/tolerance to biotic stress fac-
health-certification schemes demand free- tors. Many sites chosen for apple produc-
dom from this pathogen. Fire blight is also tion are infested with damaging pests or
damaging and rootstocks should be tested diseases. It is essential that these problems
and guaranteed free of this bacterial are recognized prior to orchard establish-
pathogen. Most countries offer schemes ment and rootstocks chosen to provide
whereby rootstocks are certified as resistance/tolerance to the problems.
• Tolerance to abiotic stress factors. Where rating clonal rootstocks for raising these
sites suffer from transient drought or compact scion types.
asphyxiation of the soil, rootstocks should Apple rootstocks are still raised from seed
be chosen to provide some tolerance of quite extensively in China, where M. sieversii
these conditions. Where severe winter and M. prunifolia are popular choices. These
cold is a problem rootstocks with cold tol- seedling types are reported to give deep
erance are essential. rooting and good drought tolerance, as well
• Freedom from suckering. Rootstocks that as tolerance to severe winter-cold conditions.
produce many suckers, either from their Japanese growers have traditionally used
shanks or from their root systems, are a seedling-raised rootstocks, although this
problem to the fruit grower. The suckers practice has begun to change in recent years
inhibit weed control practices, increase with the use of M.26 interstocks and the
chances of pest and disease infection, development of clonal dwarfing rootstocks.
compete with the tree and are expensive More recently, Chinese growers have begun
to remove annually. to use M.26 as an interstock to provide some
degree of scion growth control when using
invigorating seedling-raised rootstocks of M.
5.8 Apple Rootstocks Propagated from prunifolia or M. sieversii.
Seed
Fruit growers in very few countries now 5.9 Vegetatively Propagated Rootstock
rely on apple rootstocks raised from seed. Clones and Sub-clones Used to Control
Only in countries where clonal propagation Tree Vigour
of rootstocks has proved too difficult or
uneconomic are seedling rootstocks still Most apple-scion cultivars grown on their
used to any significant extent. Seedling root- own roots or on seedling rootstocks produce
stocks, mainly of the scion cultivar ‘Red large standard trees of 7–10 m in height and
Delicious’, were used quite extensively in spread. Whilst such trees are acceptable in
the USA until relatively recently. Their main countries where land and labour are very
use was for raising trees of compact- or inexpensive, in most apple-producing areas of
spur-type scion clones, where control of tree the world some reduction in this natural
vigour by use of a dwarfing rootstock was vigour is desired. Clonal apple rootstocks have
not a priority. More recently, this use has been available for many years, which offer a
been superseded by the use of semi-invigo- full range of scion-vigour control (Fig. 5.5).
1m
M.27 M.9 M.26 MM.106
1m
MM.111 M.25
Fig. 5.5. Silhouettes of ‘Cox’s Orange Pippin’ apple-trees grafted on a range of Malling and Malling–Merton
rootstocks, showing their effect on scion vigour control.
Rootstocks providing vigour control simi- planted in high-density planting systems on

lar to that of M.27 are often referred to as deep, highly fertile soils with adequate sup-
super dwarfing and those similar to M.9 as plies of water (Plate 5.1) Other new root-
dwarfing. Similarly, rootstocks with vigour stocks showing preliminary promise in this
similar to M.26 are classed as semi-dwarfing vigour category are AR.10-2-5, AR.628-2 and
and those similar to MM.106 as semi-vigor- AR.69-7 from the HRI-East Malling breeding
ous. The vigorous rootstocks are those simi- programme (Webster and Tobutt, 2001).
lar to M.25 or to seedlings. In recent years,
these rootstock vigour categories have
become indistinct, as many new rootstocks 5.9.2 Dwarfing selections
exhibiting vigour intermediate between the
main categories have been selected. 5.9.2.1 M.9 and its ‘sub-clones’
Nevertheless, an attempt is made below to
list and categorize some of the many root- The tree vigour currently preferred by most
stocks available for apple. apple producers is that epitomized by trees
on M.9. The final size of trees grown on M.9
will depend greatly on the inherent vigour of
5.9.1 Super dwarfing selections the scion cultivar, the soil fertility and the
management system adopted by the grower.
The most dwarfing rootstock, which has However, the aim in many parts of northern
been available from commercial nurseries for Europe is to grow trees that are no more than
many years, is the super dwarfing selection 3 m in height, so that all pruning and har-
M.27. When grafted with most scion culti- vesting can be achieved without using lad-
vars this rootstock produces trees that are 2.5 ders (Plate 5.2). Where light levels are very
m or less in height and spread. Although of good, trees on M.9 are often grown some-
great value to fruit growers wishing to estab- what taller (as in the central axe system of
lish very high-density planting systems, it is training) but, although yields are increased
unsuited to many soils and sites. Trees on using such systems, so also are the costs of
M.27 have relatively shallow root systems management if more work using ladders or
and are not suited to poor, infertile, shallow driven platforms becomes necessary.
or droughty soils. The rootstock induces Compared with many of the other popu-
heavy fruit set and, unless fruitlets are lar clonal rootstocks for apple (e.g. M.26 and
thinned severely, fruit sizes will be smaller MM.106), M.9 is rather less productive on the
than for trees on M.9. One of the disadvan- nursery layer bed. This proved a particular
tages listed above for M.27 (i.e. sensitivity to problem with M.9A, the first selection of M.9
drought and associated small fruit size) is produced free from major viruses. Although
also associated with several other rootstocks M.9 EMLA, the first selection produced free
more recently released in the same vigour of all known major and latent viruses, was
category. Trials at Horticulture Research easier than M.9A to propagate, many nurs-
International (HRI)-East Malling have shown erymen still felt that improved propagation
that trees on B.146 and P.22 are extremely was an important goal. As a consequence,
dwarfing and produce very poor fruit size nurserymen and researchers based in
when grown without supplementary irriga- Belgium, The Netherlands, Germany and
tion (Webster and Hollands, 1999a), even in France began to reselect from within layer
the relatively moist climatic conditions of the beds of M.9, with the objective of identifying
UK. However, on irrigated and highly fertile new ‘sub-clones’ of M.9 that exhibited better
soils, such as those found in The propagation characteristics. Several sub-
Netherlands, both of these rootstocks per- clones are now marketed, especially in
form better than M.27. Europe, and it is often difficult for the fruit
M.27 and other newer rootstocks of simi- producer to determine which one to choose,
lar vigour (see Table 5.1) are best used for as the evidence from trials comparing these
vigorous (e.g. triploid) scion varieties sub-clones is sometimes inconsistent.
Apples - Chap 05
104
Table 5.1. Some traditional and more recently released super dwarfing rootstocks for apples.
Rootstock Origin Parents Availability of rootstock (2001) Remarks References
M.27 HRI-East Malling, UK M.13 × M.9 Widely available Super dwarfing; induces good precocity Preston, 1954, 1971;
21/3/03
and high yield efficiency; may induce small Barritt et al., 1995
fruit size; poorly anchored, sensitive to
winter cold, drought and woolly apple aphid
B.146 Michurinsk College, Not known Unavailable for commercial Vigour variable depending upon soil type Zagaja et al., 1988;
2:55 pm
Russia plantings and irrigation (M.27 to M.26); good yield Wertheim, 1991;
efficiency; brittle roots; bad burr-knots and Barritt et al., 1995
suckering; winter-hardy; red leaves
B.491 Michurinsk College, Not known Unavailable for commercial Vigour similar to M.27; good yield efficiency; Wertheim, 1991;
Russia plantings winter-hardy; high uptake of calcium into Callesen, 1997
Page 104
leaves and fruits; easy to propagate
BM 427 Balsgård, Sweden M.4 × ‘Antonovka Limited availability for High yield efficiency; winter-hardy; otherwise Trajkovski and
Kamensischka’ commercial plantings similar to M.27; very limited trials/information Andersson, 1980
available currently
G.65 Cornell University, M.27 × ‘Beauty’ Temporarily unavailable Similar or less vigour than M.27; growth too Robinson et al.,
New York, USA crab apple weak in most situations; high yield precocity 1999
and efficiency; suckers; moderately resistant
to fire blight and collar rot
JM.1, 5 and 8 Apple Research ‘Marubakaido’ × M.9 Experimental rootstocks, Preliminary evidence from Japan suggests Bessho and
Centre, NIFTS, Japan still under evaluation vigour similar to M.27; enhanced fruit firm- Soejima, 1992;
ness and sugar content; resistant to collar Soejima et al., 1998
rot; very limited trials/information available
currently
J-TE-G Techobuzice, Czech M.9 × ‘Croncels’ Available in Europe Similar vigour to M.27; very high yield Dvorák, 1988
Republic efficiency; in other respects similar to M.27
M.20 HRI-East Malling, UK Chance seedling Limited availability in the UK, Similar vigour to M.27; good yield precocity Jackson, 1986;
The Netherlands and Canada and efficiency; induces better fruit size than Wertheim, 1992
M.27; poor anchorage; suckers more than
M.27; difficult to propagate
Apples - Chap 05
P.22 (‘Last Skierniewice, Poland M.9 × ‘Antonovka’ Widely available Much more invigorating than M.27 if planted Zagaja, 1980; Barritt
Minute’) on fertile soils with adequate supplies of et al., 1995;
water; on drier soils tree size very much Kruczynska and
reduced; three sub-clones – J and K (more Czynczyk, 1998;
21/3/03
juvenile) and S (more adult) – available in Webster and
Europe; high yield precocity and efficiency; Hollands, 1999a;
anchorage similar to M.27, some suckering; Czynczyk and
winter-hardy; sensitive to drought and to Piskor, 2000
2:55 pm
woolly apple aphid; tolerant to collar rot
P.59 (‘Polan Skierniewice, Poland A.2 × B.9 Limited availability in Europe Similar vigour to P.22 in Polish trials; good Jakubowski,
59’) yield efficiency; red leaves; very limited trials/ 1999a,b
information available currently
P.16 × M.26
Page 105
P.61 Skierniewice, Poland Unavailable for commercial Very dwarfing, less than P.22 in Polish trials; Jakubowski,
plantings very high yield efficiency; very limited trials/ 1999a,b
P.66 Skierniewice, Poland P.22 × M.26 Unavailable for commercial Vigour greater than P.61 but significantly less Jakubowski,
Apple Rootstocks
plantings than M.9; good yield efficiency; very limited 1999a,b
trials/information available currently
V.3 Vineland, Canada ‘Kerr’ crab apple Limited availability for trials in Variable vigour between P.22 and M.9; high Elfving et al., 1993;
open-pollinated the USA and Europe yield efficiency; winter-hardy; limited trials Barritt et al., 1995
information to date
Voinesti 2 Voinesti, Romania M.9 × ‘Cretesc’ Currently available only in Vigour similar to M.27 or P.22 in Dutch trials; Parnia et al., 1997
country of origin good yield efficiency; very limited trials/
NIFTS, National Institute of Fruit Tree Science.
105
It is generally accepted that most of tion in the orchard have been studied in
these new sub-clones of M.9 are easier to several countries and the results are slightly
propagate than the traditional virus-free variable (Wertheim, 1997; Webster and
M.9 EMLA sub-clone. Although all are sim- Hollands, 1999b). Small differences in tree
ilar genetically, there are ontogenetic dif- vigour are noted, with the Dutch sub-clone
ferences between the sub-clones. The more Fleuren 56 currently the most dwarfing and
adventitious (often wrongly called juve- the Belgian sub-clone K (Nicolai) 29 among
nile) types usually have slightly narrower the more invigorating. However, the differ-
leaves, branch more freely and are easier to ences between the most dwarfing and invig-
propagate, whereas the less adventitious orating clones are rarely more than 15%.
(or more adult) types show the opposite Trials in the UK testing most of the sub-
characteristics (Van Oosten, 1986). Trials at clones have shown no significant differ-
East Malling conducted some years ago ences in their effects on scion yield
(Webster and Jones, 1989) showed that precocity and efficiency if trees of similar
propagation of the EMLA sub-clone of M.9 size and branching are planted (Webster
could also be improved if it was put and Hollands, 1999b). Growers wishing to
through an extended in vitro phase prior to use M.9 should choose the more invigorat-
establishing hedges or stool beds. ing sub-clones, where the site and soils are
Repeated subculturing in micropropaga- slightly suboptimal or where a weak scion
tion induced a form of adventitiousness or cultivar is chosen and planted at medium
false juvenility, which persisted for many tree densities. Where soils are highly fertile,
years after establishment in the nursery if scions have strong inherent vigour and very
the hedges/stools were severely pruned high-density planting systems are chosen,
each year. the weaker sub-clones should be chosen.
The comparative attributes of the various Table 5.2 lists a few of the more popular
M.9 sub-clones when used for fruit produc- sub-clones of M.9.
Table 5.2. ‘Sub-clones’ of M.9 available as virus-free rootstocks.
Name or number of sub-clone Origin Remarks
M.9 A UK The first sub-clone freed of major viruses; a poor

clone in the nursery
M.9 EMLA UK The first M.9 sub-clone freed of all known major
and latent viruses; moderate vigour; less easy to
propagate than many sub-clones
Burgmer 719 (= B.1) Germany Slightly more invigorating than the EMLA sub-
clone in UK trials
clone in UK trials
clone in UK trials
Fleuren 56 The Netherlands Selected by the Fleuren nursery; the least
vigorous sub-clone known currently
NAKB T.337 The Netherlands The most often used of the four virus-free sub-
clones produced by NAKB
K (Nicolai) 29 (RN 29) Belgium One of the more invigorating ‘sub-clones’ of M.9
Pajam 1 (Lancep) France Easier to propagate and slightly less vigorous than
M.9-EMLA
Pajam 2 (Cepiland) France Similar or slightly more vigorous than M.9-EMLA;
easier to propagate
5.9.2.2 Alternatives to M.9 as dwarfing 5.9.3 Semi-dwarfing selections

rootstocks
Where climatic and soil conditions are
Breeders of apple rootstocks have tried, over
unsuited to use of M.9, apple growers requir-
the last 40 years, to produce rootstocks with
ing dwarfed trees often turn to slightly more
similar vigour and attributes to M.9 but
invigorating clonal rootstocks such as M.26 or
which have additional advantages.
other rootstocks of similar vigour potential,
Particular goals have been improved resis-
which induce 15–30% more vigour and tree
tance to winter-cold damage (see Russian,
size than M.9 (Plate 5.3). M.26 exhibits much
Polish and Canadian rootstock selections)
better tolerance of winter cold than M.9
and to fire blight (see Geneva, New York
(Ferree and Carlson, 1987) and for this reason
State, rootstock selections). Although several
has often been chosen for areas where winters
new rootstocks exhibiting increased resis-
tances to these and other problems have are occasionally severe. It is also frequently
been produced, some of them are, unfortu- used as a dwarfing interstock (see below) in
nately, slightly inferior to M.9 in one or more countries such as Japan and China. Like M.9,
other characteristics (e.g. yield efficiency, trees on M.26 need stakes or other supports in
drought tolerance, fruit size, etc.). all but the most sheltered locations.
Mark (previously MAC.9), a rootstock Although a useful rootstock when used in
bred at Michigan State University in the appropriate situations, M.26 does have sev-
USA, showed great promise initially as a eral disadvantages. It has a tendency to pro-
rootstock with vigour between M.9 and M.26 duce many burr-knots on the above-ground
that induced excellent yield precocity and rootstock stem (shank) and these can prove to
efficiency (Carlson and Perry, 1986; Perry, be sites for entry of damaging pests and
1990). Unfortunately, it has performed very pathogens. Excessive burr-knotting also
poorly in some trials, especially those reduces growth on the scion and this can
planted on hot dry soils. It is very sensitive result in uneven growth in orchards planted
to drought (Fernandez et al., 1997) and to on M.26. This is especially severe where trees
crown gall and develops a swelling at or just are machine-planted with differing amounts
below ground level, which reduces tree per- of rootstock shank exposed above ground
formance on some sites (Stover and Walsh, level. Growers choosing M.26 should plant
1994) and increases tree-to-tree variability in the trees with their graft unions as close as
the orchard. possible to the soil surface, without causing
Most of the other rootstocks produced scion rooting; this should greatly reduce the
and released with vigour similar to M.9 problems of burr-knotting and uneven tree
have been bred to provide improved toler- growth. Although sub-clones of M.26 exhibit-
ance to winter-cold injury and should be ing less burr-knotting have been selected in
chosen in preference to M.9 in areas where The Netherlands (Denissen et al., 1993;
very low winter temperatures are common. Wertheim and Kunneman, 1993), these have
However, as can be noted in Table 5.3, yet to gain widespread popularity elsewhere.
some of these selections are inferior to M.9 There are also a few reports of scions on
in one or more of their other attributes and M.26 rootstock being less efficient in calcium
care must be taken when choosing among uptake than trees on M.9 and other rootstocks.
them. Like M.9, these rootstocks generally Whilst unlikely to be a problem with cultivars
require staking. efficient in calcium uptake (e.g. ‘Gala’), M.26 is
Other rootstocks in this vigour category not the best choice for cultivars such as ‘Cox’s
that are showing initial promise in experi- Orange Pippin’, where calcium-related disor-
mental trials are AR.680-2, AR.295-6 and ders, such as bitter pit, can prove problematic.
AR.486-1 from the HRI-East Malling pro- M.26 is also very sensitive to fire blight and
gramme and G.16 and CG41 from the fruit breeders based at Cornell University in
Cornell Geneva programme. the USA are currently developing transgenic
Apples - Chap 05
108
Table 5.3. Some of the traditional and new dwarfing rootstock selections for apples.
Rootstock
name or
21/3/03
number Origin Parents (if known) Availability (2001) Remarks References
M.9 (‘Jaune Reselected at HRI- Chance seedling Widely available The most popular dwarfing rootstock; induces Hatton, 1917; Van
de Metz’) East Malling, UK found in France excellent yield precocity and efficiency; induces Oosten, 1977, 1986;
large fruit size; brittle roots; poor anchorage; Webster and
2:55 pm
some suckering, depending upon scion cultivar, Hollands, 1999b
rootstock sub-clone and site conditions; sensitive
to winter cold, poor drainage and shows some
drought sensitivity; sensitive to fire blight and to
woolly apple aphid; some resistance to collar rot
Page 108
B.9 Michurinsk College, M.8 × ‘Red Standard’ Available Vigour between M.9 and M.26; good yield Barritt et al., 1995;
Russia efficiency; anchorage slightly better than M.9; Kruczynska and
winter-hardy; sensitive to woolly apple aphid; Czynczyk, 1998;
resistant to collar rot; propagation difficult; red Webster and
leaves; exhibited more field tolerance to fire blight Hollands, 1999a
infections than M.9
B.469 Michurinsk College, Not known Experimental rootstock, Vigour similar to M.9; induces good yield and fruit Wertheim, 1991
Russia still under evaluation size; winter-hardy; very limited trials/information
available currently
C.6 Louisiana, USA M.8 open-pollinated Unavailable Variable vigour M.9 to M.26; precocity, yield Barritt et al., 1995
commercially in Europe efficiency and fruit size variable but mainly similar
to M.9; very limited trials/information available
currently
G.16 Cornell University, Ottawa 3 × Malus Limited availability in Similar vigour to M.9; good yield efficiency; Johnson, 1999;
New York, USA floribunda USA and Europe resistant to fire blight and collar rot; very limited Robinson et al.,
trials/information available currently 1999
JM.2 Apple Research ‘Marubakaido’ × M.9 Experimental Vigour similar to M.9 in Japanese trials; tolerant Bessho and
Centre, NIFTS, rootstocks, still under of woolly apple aphid; resistant to collar rot; easy Soejima, 1992
Japan evaluation propagation from hardwood cuttings; very limited
JM.7 Apple Research ‘Marubakaido’ × M.9 Experimental Vigour similar to M.9 in Japanese trials; tolerant of Bessho and
Centre, NIFTS, rootstocks, still under woolly apple aphid; resistant to collar rot; easy Soejima, 1992
Japan evaluation propagation from hardwood cuttings; very limited
Apples - Chap 05
J-OH-A Omomoue-Holice, Not known Limited availability in Vigour slightly greater than M.9; high yield Dvorak, 1983;
Czech Republic Europe efficiency; anchorage similar to M.9; possible virus Mantinger, 1996
sensitivity; very limited trials/information available
currently
21/3/03
Jork (J).9 Jork Institute, M.9 open-pollinated Available in Europe Vigour similar to or slightly more than M.9; good Faby et al., 1986;
Germany yield efficiency and fruit size; some burr-knots and Wertheim, 1991
suckers; more hardy than M.9; tolerates soil/
environmental stresses well; resistant to collar rot;
easy to propagate
2:55 pm
J-TE-E Techobuzice, Not known Limited availability in Vigour similar to M.9; high yield efficiency; Dvorák, 1988
Czech Republic Europe anchorage similar to M.9; possible virus sensitivity;
very limited trials/information available currently
J-TE-F Techobuzice, Not known Limited availability in Vigour similar to M.9; medium to high yield Dvorák, 1988
Czech Republic Europe efficiency; anchorage similar to M.9; possible virus
Page 109
sensitivity; very limited trials/information available
currently
M.8 (‘Clark Reselected at HRI- Originally known as Available in genetic Vigour similar to M.9; good yield efficiency; very Hatton, 1917; Brase,
Apple Rootstocks
Dwarf’) East Malling, UK ‘French Paradise’ collections poor anchorage due to brittle roots; suckers more 1954
than M.9; sensitive to poor drainage and drought;
sensitive to woolly apple aphid
MAC.9 Michigan State M.9 open-pollinated Limited availability Vigour variable depending upon soil type; usually Carlson, 1980; Perry,
(‘Mark’) University, USA between M.9 and M.26; very good yield efficiency; 1990; Barritt et al.,
anchorage slightly better than M.9; swellings at 1995; Webster and
soil line; drought-sensitive and poor performance Hollands, 1999a
in hot dry soils; more resistant to collar rot than
M.9; weak union, with some triploid scions
MAC. 39 Michigan State M.9 open-pollinated Not available Vigour slightly less than M.9; suckers badly; very Carlson, 1980;
University, USA commercially limited trials/information available currently Barritt et al., 1995
Ottawa 3 Ottawa, Canada ‘Robin’ × M.9 Limited availability Vigour similar to M.9; good yield productivity; Granger, 1984;
anchorage similar to M.9; winter-cold-tolerant; Quamme and
very sensitive to virus; resistant to collar rot; Brownlee, 1990;
sensitive to woolly apple aphid; very difficult to Barritt et al., 1995
propagate by conventional techniques
P.2 (‘Skilig’) Skierniewice, Poland M.9 × ‘Antonovka’ Available in Europe On fertile soils vigour similar to M.9; but it suffers Zagaja, 1980; Barritt
from transient drought; tree size is much smaller; et al., 1995;
good yield productivity; fruit size poor unless Kruczynska and
irrigated sufficiently; anchorage similar to M.9; Czynczyk, 1998;
suckers slightly; winter-hardy; sensitive to woolly Webster and
109
aphid; resistant to collar rot Hollands, 1999a
Continued
Apples - Chap 05
110
Rootstock
name or
21/3/03
P.16 (‘Lizzy’) Skierniewice, Poland ‘Longfield’ × M.11 Available Vigour very variable, depending upon soil Zagaja, 1980; Barritt
conditions; on fertile soils equal to M.9, on poorer et al., 1995;
soils similar to M.27; very yield-efficient; fruit size Kruczynska and
2:55 pm
usually less than on M.9; anchorage similar to M.9; Czynczyk, 1998;
may sucker profusely on some sites; hardiness Webster and
similar to M.9; sensitive to woolly aphid; resistant Hollands, 1999a
to collar rot
P.60 Skierniewice, Poland A.2 × B.9 Available in Europe Similar vigour to P.2; good yield efficiency; suffers Zagaja et al., 1991;
Page 110
(‘Polan 60’) from burr-knotting and superficial rooting; winter- Jakubowski,
hardy; red leaves; very limited trials/information 1999a,b; Kurlus and
available currently Ugolik, 1999
P.62 Skierniewice, Poland A.2 × M.27 Currently available Vigour slightly less than for M.9 in Polish trials; Jakubowski,
only for trials very high yield efficiency; very limited trials/ 1999a,b
P.67 Skierniewice, Poland A.2 × P.2 Currently available Vigour similar to M.9 in Polish trials; very high Jakubowski,
only for trials yield efficiency; very limited trials/information 1999a,b
available currently
‘Supporter 1’ Dresden, Germany M.9 × Malus baccata Increasing availability Vigour 20% less than M.9; high yield efficiency; Fischer, 1997, 1999
similar hardiness to M.9; as yet minimal evidence
available from trials in countries other than
Germany
‘Supporter 2’ Dresden, Germany M.9 × Malus × Increasing availability Vigour 15% less vigorous than M.9; high yield Fischer, 1997, 1999
micromalus efficiency; hardy; almost no evidence yet available
from trials in countries other than Germany
‘Supporter 3’ Dresden, Germany M.9 × Malus × Increasing availability Vigour similar to M.9; high yield efficiency; hardy; Fischer, 1997, 1999
micromalus almost no evidence yet available from trials
in countries other than Germany
V.1 Vineland, Canada ‘Kerr’ crab apple Experimental; still Slightly more invigorating than M.9; similar yield Elfving et al., 1993;
open-pollinated under evaluation efficiency; winter-hardy; very limited trials/ Barritt et al., 1995
NIFTS, National Institute of Fruit Tree Science.

sub-clones of M.26 that have genes introduced Malling programme have performed well in
that are aimed at providing increased resis- trials in the UK and New Zealand. AR.86-1-
tance to this damaging pathogen. 25 (soon to be named M.116) and AR.86-1-20
Fruit breeders have endeavoured to pro- have produced trees similar in growth and
duce new rootstock clones with similar yields to trees on MM.106, but with much
vigour and cold tolerance to M.26 but with improved resistance to collar rot. Some of the
no burr-knotting and improved induction of traditional and/or currently popular semi-
yield precocity and efficiency. A few of these invigorating and invigorating apple root-
are listed in Table 5.4. stocks are listed and described in Table 5.5.
Other experimental rootstocks showing
promise in this vigour category but which
have yet received only limited evaluation are 5.9.5 Very vigorous selections
AR.801-11 from the HRI-East Malling pro-
gramme and several selections from the There is nowadays little demand for vigor-
Geneva programme. ous clonal rootstocks for apple cultivation.
Many of the early selections, such as M.1,
M.2, M.12, M.16 and Crab C, have disap-
5.9.4 Semi-vigorous to vigorous selections peared from commerce. Only Merton 793
remains a popular rootstock, especially in
Although most orchards of 50 or more years the southern hemisphere. Table 5.6 lists a few
ago were planted on semi-vigorous or vigor- of the invigorating rootstock clones.
ous rootstocks, these have become much less
popular in recent times (Plate 5.4). They are
still used for dessert-apple plantings on very 5.10 Clonal Rootstocks Used to Adapt
poor soils where dwarfing rootstocks will not Trees to Unfavourable Environmental
thrive. Also, they are still very popular with Conditions
producers of apples grown for the apple-juice
or cider markets. Cider-apple trees are har- 5.10.1 Tolerance to winter-cold injury
vested using mechanical shakers and the
fruits are mechanically collected up from the Several of the traditional and most popular
orchard floor. Most dwarfing rootstocks are rootstocks, such as M.9 and M.27, exhibit poor
unsuited to mechanical shaking, on account tolerance to severe winter cold. In some situa-
of their shallow and often brittle root systems. tions, especially on poorly drained soils, trees
For many years the two most popular on sensitive rootstocks may be killed in severe
semi-dwarfing rootstocks have been M.7 and winters. It should be stressed that many
MM.106. M.7 is popular in the USA on dwarfing rootstocks thrive best when the soil
account of its good adaptability to different is well drained, either naturally or artificially.
soil types, its induction of good precocity Where there is inadequate snow cover, cou-
and yield efficiency and its resistance to col- pled with very low temperatures, much dam-
lar rot and fire blight. However, it suckers age is caused to the roots and shanks of
and is susceptible to winter injury and trials sensitive rootstocks. After severe winter frosts,
in Europe have shown it to be less produc- damage can frequently be observed as death
tive than MM.106. This latter stock induces of cambial tissues in the rootstock shank.
excellent yield precocity and productivity Rootstock breeders centred in Russia,
but is sensitive to collar and crown rots as Poland, the Ukraine, Belarus, Canada and the
well as fire blight. Recently, several root- USA have made tolerance to low winter tem-
stocks have been tested from within the peratures a principal goal in their pro-
Cornell Geneva rootstock-breeding programmes. However, care must be taken in
gramme that have shown early promise. choosing specific rootstocks from these pro-
Several exhibit resistance to fire blight, collar grammes, as not all the selections exhibit cold
rot and woolly apple aphid. In addition sev- tolerance. For instance, the Polish selections
eral rootstocks selected from the HRI-East P.1 and P.16 show similar sensitivity to M.9.
Apples - Chap 05
112
Table 5.4. Some of the traditional and recently released semi-dwarfing rootstocks for apples.
Rootstock
name or
21/3/03
M.26 HRI-East Malling, UK M.16 × M.9 Widely available More vigorous than M.9, but less vigorous than Preston, 1954,
MM.106; average yield efficiency; good fruit size 1970; Rogers,
and colour; anchorage better than for M.9 but 1958; Proctor et
2:55 pm
needs staking on most exposed sites; most sub- al., 1974
clones burr-knot badly; winter-hardy; average
drought tolerance; sensitive to woolly apple aphid
and fire blight; poor calcium uptake on some sites;
poor tolerance to heavy/wet soils; relatively easy
Page 112
to propagate
B.62-396 Michurinsk College, Not known Available for trials in Vigour variable in trials in the Ukraine and Russia; Hulko and Hulko, 1999;
Russia several countries in usually between M.9 and M.26; very hardy; very Kapichnikova, 1999;
eastern and central limited trials/information available currently Kuldoshin, 1999;
Europe Kurlus and Ugolik,
1999; Verzilin et al.,
1999
‘Bemali’ Balsgård, Sweden ‘Mank’s Codlin’ × M.4 Limited availability Vigour greater than M.9 and more similar to M.26; Trajkovski and
much less yield-efficient than M.9; produces small Andersson, 1980
fruit size; better winter-hardiness than M.9;
resistant to fire blight
G.11 Cornell University, M.26 × Robusta 5 Increasing availability Similar vigour to M.26; similar or better yield Johnson, 1999;
New York, USA efficiency than M.26; suckers; resistant to collar Robinson et al., 1999
rot and to fire blight; very limited trials/information
available currently in Europe
J-TE-H Techobuzice, M.9 × ‘Croncels’ Limited availability Similar or slightly greater vigour than M.26; poor Dvorák, 1988
Czech Republic yield efficiency in some trials; very limited trials/
KSC 28 Kentville, Nova ‘Beautiful Arcade’ × Available in Canada Vigour greater than M.26; yield efficiency variable; Privé and Embree,
Scotia, Canada ‘Antonovka winter-hardy 1997
Kamensischka’
P.1 Skierniewice, Poland M.4 × ‘Antonovka’ Available in Poland More invigorating than M.26; poor yield precocity Zagaja, 1980; Barritt et
and efficiency; severe burr-knotting; sensitive to al., 1995; Kruczynska
viruses and Czynczyk, 1998
Apples - Chap 05
P.14 Skierniewice, Poland M.9 open-pollinated Available in Europe Vigour similar or greater than on M.26; good yield Czynczyk and
(‘Skidal’) efficiency in Polish trials; poor precocity in Olszewska, 1990;
Hungarian trials; very limited trials/information Kurlus and Ugolik, 1999
available currently
21/3/03
‘Supporter 4’ Dresden, Germany M.9 × M.4 Available Variable yield efficiency compared with M.26; Fischer, 1997, 1999
(‘Pillnitz 80’) hardy; easy to propagate
V.7 Vineland, Canada ‘Kerr’ crab apple Experimental root- Vigour slightly greater than M.26; similar yield Elfving et al., 1993;
open-pollinated stock still under efficiency; winter-hardy; very limited trials/ Barritt et al., 1995
2:55 pm
evaluation information available currently
Page 113
Apple Rootstocks
113
Apples - Chap 05
114
Table 5.5. Some of the traditional and more recently developed semi-invigorating/invigorating rootstocks for apples.
Rootstock
name or
21/3/03
M.7 East Malling, UK Not known Widely available Similar to or slightly more invigorating than MM. Hatton, 1917;
106; induces average yield productivity; Preston, 1970
suckering often a problem; sensitive to winter-
2:55 pm
cold injury; field-tolerant to collar rot and fire
blight; good adaptability to soil types
MM.106 East Malling, UK ‘Northern Spy’ × M.1 Widely available High yield efficiency; fruit size can be smaller than Preston, 1955, 1966;
on M.9; good anchorage; few suckers; average Parry, 1965
Page 114
tolerance of winter cold; some drought tolerance;

susceptible to collar and crown rot, fire blight and
tomato ringspot virus; resistant to woolly apple
aphid
MM.111 East Malling, UK ‘Northern Spy’ × Widely available Slightly more invigorating than MM.106; poorer Preston, 1955, 1966;
Merton 793 precocity but good yield efficiency when trees Parry, 1965
mature; good anchorage; few suckers; some
tolerance of winter cold; sometimes sensitive to
collar and crown rots; resistant to woolly apple
aphid
G.30 Geneva Research Robusta 5 × M.9 Increasing availability Similar vigour to M.7 in US trials; more yield- Johnson, 1999;
Station, New York, efficient and precocious than M.7; suckers; Robinson et al.,
USA resistant to fire blight; resistant to crown rot; 1999
susceptible to woolly apple aphid; weak graft
unions a problem; very limited trials/
information available currently outside USA
G.210 Geneva Research Ottawa 3 × Robusta 5 Experimental rootstock Similar vigour to M.7; yield efficiency similar to Johnson, 1999;
Station, New York, still under evaluation M.26 and better than M.7; suckers; resistant to Robinson et al.,
USA collar/crown rot, woolly apple aphid and fire 1999
blight; very limited trials/information available
currently outside USA
KSC.7 Kentville, Canada ‘Beautiful Arcade’ × Limited availability in Semi-vigorous to vigorous; average to poor yield Privé and Embree,
‘Antonovka’ Canada efficiency; winter-hardy; very limited trials/ 1997
Apples - Chap 05
KSC.11 Kentville, Canada ‘Beautiful Arcade’ × Limited availability in Semi-vigorous to vigorous; average to poor Privé and Embree,
‘Antonovka’ Canada yield efficiency; very limited trials/information 1997
available currently
KSC.24 Kentville, Canada ‘Beautiful Arcade’ × Limited availability in Semi-vigorous to vigorous; average to poor Privé and Embree,
21/3/03
‘Antonovka’ Canada yield efficiency; very limited trials/information 1997
available currently
M.4 East Malling , UK Originally ‘Holstein Limited availability in Similar vigour to M.7; induces variable yield Hatton, 1917;
Doucin’ genetic collections efficiency; poor anchorage; resistant to fire blight Preston, 1970
2:55 pm
and to collar rot; poor uptake of potassium
M.116 HRI-East Malling, UK MM.106 × M.27 Experimental rootstock Induces vigour and cropping similar to MM.106; Webster et al., 1986
(AR.86-1-25) still under evaluation very resistant to collar/crown rot and to woolly
apple aphid; only limited trials data currently
available from UK and New Zealand
Page 115
MM.104 East Malling, UK M.2 × ‘Northern Spy’ Limited availability Vigour slightly greater than MM.106; average Preston, 1955;
precocity and productivity; tolerates dry soils; Parry, 1965
sensitive to collar and crown rots; resistant to
Apple Rootstocks
woolly apple aphid and fire blight
V.2 Vineland, Canada ‘Kerr’ crab apple Experimental rootstock More invigorating than M.26 but less than P.1; Elfving et al., 1993;
open-pollinated still under evaluation good yield efficiency; winter-hardy; only limited Barritt et al., 1995
information available from trials in countries other
than the USA and Canada
115
Apples - Chap 05
116
Table 5.6. Some traditional and more recently released invigorating rootstocks for apples.
Rootstock
name or
21/3/03
Merton (MI) John Innes Institute, M.2 × ‘Northern Spy’ Widely available in the Semi-vigorous to vigorous; induces poor yield
793 UK southern hemisphere precocity but average yield efficiency when trees
mature; good anchorage; few suckers; resistant
2:55 pm
to woolly apple aphid and crown rot; very suitable
for replant situations on poor soils
‘Alnarp 2’ Alnarp Research Not known Available Very invigorating; poor yield efficiency; good Pieniazek et al.,
Station, Sweden anchorage and winter-cold tolerance 1976
B.54-118 Michurinsk College, Not known Available in Russia and Greater vigour than MM.106; variable yield Hulko and Hulko,
Page 116
Russia some other parts of efficiency; good anchorage; winter-hardy; good 1999; Kuldoshin,
eastern Europe drought tolerance; resistant to fire blight 1999
B.57-490 Michurinsk College, Not known Available in Russia and Vigorous; poor precocity and productivity in Kurlus and Ugolik,
Russia some other parts of Hungarian and Dutch trials; winter-hardy; 1999
eastern Europe resistant to fire blight; very limited trials/
KSC.3 Kentville, Nova ‘Beautiful Arcade’ × Limited availability in Very vigorous; yield efficiency good; good Privé and Embree,
Scotia, Canada ‘Antonovka’ Canada anchorage; winter-hardy 1997
KSC.6 Kentville, Nova ‘Beautiful Arcade’ × Limited availability in Very vigorous; average yield efficiency; good Privé and Embree,
Scotia, Canada ‘Antonovka’ Canada anchorage; winter-hardy 1997
M.1 East Malling, UK Originally named Available in genetic Vigorous tree that exhibits poor yield efficiency; Hatton, 1917
reselection ‘Broadleaved collections good anchorage, few suckers; susceptible to
Paradise’ drought
M.2 East Malling, UK Originally named Available in genetic Vigour similar to M.1; variable yield efficiency; Hatton, 1917
reselection ‘Doucin’ collections only average fruit size; good anchorage; suckers;
adapts to most soil types; some sensitivity to
drought; tolerates wet soils; moderately resistant
to fire blight; poor potassium uptake
M.25 East Malling, UK ‘Northern Spy’ × M.2 Limited availability Vigour slightly greater than MM.111; high yield Preston, 1955, 1966;
efficiency and precocity; good anchorage; few Parry, 1965
suckers; sensitive to woolly apple aphid
‘Marubakaido’ Japan Malus prunifolia Available in Japan Very vigorous; only average induction of yield
‘Ringo’ precocity and efficiency; resistant to woolly apple
aphid and collar/crown rots
Apples - Chap 05
MM.109 John Innes and M.2 × ‘Northern Spy’ Limited availability Very vigorous; only average yield precocity and Parry, 1965;
East Malling, UK efficiency; poorly anchored; performs well on Preston, 1966
droughty soils; sensitive to collar and crown rot
‘Novole’ Geneva Research M. prunifolia × Limited availability in Very vigorous; unpalatable to voles; resistant to Ferree and
21/3/03
PI286613 Station, New York, M. sieboldii the USA apple scab, fire blight and tomato ringspot virus; Carlson, 1987;
USA variable resistance to crown rot; easily Wertheim, 1998
propagated from softwood or hardwood cuttings
but difficult by layering; minimal suckering
2:55 pm
V.4 Vineland, Canada ‘Kerr’ crab apple Experimental rootstock Vigorous; average yield efficiency; winter-hardy; Elfving et al., 1993;
open-pollinated still under evaluation very limited trials/information available currently Barritt et al., 1995
Page 117
Apple Rootstocks
117
For a full review of breeding for cold tol- symptoms show until several years later, fol-
erance in apple rootstocks the reader should lowing planting of young trees in the orchard.
consult Hulko and Hulko (1999). Where Important though these pathogens are,
known, the cold tolerances of many of the they are minor compared with the damage
traditional and new clonal rootstocks are caused by fire blight in some apple-produc-
mentioned in the Remarks sections of Tables ing regions of the world (Ferree et al., 1983;
5.1 and 5.3–5.6. Perry, 1992). Rootstocks that are not only
resistant to fire blight but that also confer
some of this resistance to scions budded or
5.10.2 Tolerance to soil-borne or aerial grafted would be of great value in the east-
pathogens ern states of the USA. Amongst the currently
available rootstocks exhibiting some resis-
The roots and below-ground parts of apple tance to fire blight are G.65, G.16, G.11, G.30,
rootstocks may be attacked by many fungi G.210, Bemali, M.7, M.4, M.2, MM.104, B.118
and bacteria and the most damaging of these and B.490 (see Tables 5.1 and 5.3–5.6).
may result in tree death. The most common
species of fungi causing damage to the root-
stocks are the collar or crown rots 5.10.3 Tolerance to soil-borne or aerial pests
(Phytophthora sp.) and several studies have
investigated rootstock sensitivity to these The most damaging pest affecting apple
pathogens (Lemoine and Gaudin, 1991). rootstocks is the woolly apple aphid (E.
Among the super dwarfing rootstocks, P.22, lanigerum), which can cause significant dam-
G.65, JM.1, JM.5 and JM.8 are reported to age to roots of apple trees if they are planted
show good resistance, while M.9, Ottawa 3, on susceptible rootstocks in apple-producing
P.2, P.16, G.16, B.9, Mark and J.9 all show areas where the pest is present and where
resistance in the dwarfing category. Other soil conditions favour it. Hot and dry soils,
more invigorating rootstocks showing resis- such as those commonly used for apple pro-
tance are G.11, G.30, G.210, M.116, M.7 and duction in the southern hemisphere, are par-
‘Marubakaido’ (see Tables 5.1 and 5.3–5.6). ticularly prone to damage from woolly apple
Less common but very damaging are the aphids, although less severe damage is also
honey fungus (Armillaria mellea sensu stricto noted in parts of southern Europe and in
Vahl. ex Fr.) and other species classed in the Israel. Most dwarfing rootstocks are sensitive
USA as the southern root rots. Unfortunately, to the pest, although recent reports suggest
few, if any, apple rootstocks show resistance that several rootstocks bred in Japan (JM
to these root rots. The bacterial pathogen series) may combine dwarfing with resis-
crown gall also attacks apple rootstocks and tance. Merton 793 and the MM series of root-
may stunt growth severely in some soil con- stocks, all bred using the resistant scion
ditions. Although no rootstocks are fully cultivar ‘Northern Spy’ as one parent, exhibit
resistant to crown gall, clones do differ tolerance to the pest, as does the Japanese
slightly in their sensitivity. selection ‘Marubakaido’. Among the more
Aerial pathogens such as apple scab recently selected apple rootstocks, M.116 and
(Venturia inaequalis (Cke.) Wint.) and apple G.210 both show tolerance (see Tables 5.1
powdery mildew (Podosphaera leucotricha (Ell. and 5.3–5.6).
& Ev.) Salmon) attack rootstocks in the same Soil-borne nematodes can also cause sig-
way that they attack apple scions and protec- nificant damage to apple rootstocks and this
tive spray programmes must be applied in the is mainly a problem when planting trees on
nursery to prevent damage. Apple canker shallow, dry and acidic soils. Fortunately, the
(Nectria galligena Bres.) may also prove a prob- main apple-producing regions of the world
lem in some countries and it has been sug- only occasionally experience severe prob-
gested that this pathogen may, in some lems with nematode damage. However, the
circumstances, be carried in rootstocks and nematode species Pratylenchus penetrans
young trees produced in the nursery. Often no (Cobb) Sher & Allen has been reported to
cause damage and poor tree establishment

on sandy soils in The Netherlands.
Scion
5.10.4 Tolerance to drought or soil Graft union
asphyxiation
Interstock or interstem
Supplies of water for irrigation of apple
orchards are often limited, expensive or
both. Use of trickle irrigation delivery sys- Rootstock
tems can significantly improve the effi-
ciency of water use and is recommended for
many areas rather than the less efficient
overhead or flood irrigation methods. Fig. 5.6. Dwarfing interstock/interstem in relation to
apple scion and rootstock.
Nevertheless, rootstocks that aid efficient
water use by the tree will become more
important in the future. graft compatibility between the rootstock
Unfortunately, most dwarfing rootstocks and scion was a problem. This is a common
require some additional water supplied via practice when raising certain cultivars of
irrigation if they are to grow and crop well in pear on quince rootstocks. Interstock use in
areas experiencing very hot, dry summers. apple is less widespread and is generally
Growers choosing organic systems of apple only applicable when either the desired
production cannot use chemical herbicides dwarfing rootstock is difficult to propagate
and are often obliged to let weeds grow for on its own roots (e.g. Ottawa 3) or the soils
longer periods than are ideal beneath their are unsuited to planting dwarfing rootstocks
trees. Weeds and grass compete strongly for (e.g. infested with woolly apple aphid or
water, and tree growth on dwarfing root- subject to drought or waterlogging). Trunk
stocks can be severely impaired by excessive builders or ‘staddles’ of winter-hardy culti-
weed growth. vars, such as ‘Hibernal’, have been used for a
On stony, well-drained soils, water reten- very long time in central Europe to prevent
tion within the profile is poor and, unless the freeze damage to the tree trunk, and cold-
tree’s root system is able to penetrate deeper tolerance benefits have also been recorded
into the profile to tap water reserves, apple when using interstems of more usual length
trees will suffer drought stress. (Wertheim, 1985).
In these and several other situations, It is fortunate that, for reasons not
rootstocks exhibiting tolerance to transient understood, a significant part of the dwarf-
drought conditions can be a great aid to ing influence of an apple rootstock is attrib-
successful apple production. As a general utable to factors associated with its shank
rule, the more invigorating the rootstock, (stem piece) rather than its root system.
the less likely it is to suffer from drought. A This means that, by inserting a short length
few of the rootstocks exhibiting some of the dwarfing rootstock as an interstock
drought tolerance are described in Tables between a more invigorating rootstock and
5.1 and 5.3–5.6. the scion, the grower can achieve semi-
dwarfed trees. Up to approximately 35 cm
in length, the longer the interstock stem
5.11 Use of Interstocks and Interstems piece, the more dwarfing the effect (Parry
and Rogers, 1972). Trials in Poland have
Interstock or interstem trees are comprised shown increased productivity when using
of three genetically different components P.2 as an interstock (Kruczynska and
(Fig. 5.6). Czynczyk, 1998).
Traditionally, interstocks (interstems) Trees raised with interstocks are slightly
were used in raising fruit trees only when more expensive to produce but often their
benefits warrant this extra expenditure by the precocity of cropping and winter-
the grower. Further information on raising hardiness of the tree. Interstocks can also
trees with interstocks (interstems) can be help regulate the vigour of shoot growth on
found in Chapter 6. the scions (Wertheim and Callesen, 2001).
Occasionally, stem pieces of other scion Also, in The Netherlands an interstem of
cultivars (such as ‘Golden Delicious’ or ‘Dubbele Zoete Aagt’ is used with the scion
‘Summerred’) are inserted between the root- cultivar ‘Cox’s Orange Pippin’ to prevent
stock and the chosen scion. This can improve trunk rot caused by P. cactorum.
Further General Reading on Apple Rootstocks
Ferree, D.C. and Carlson, R.F. (1987) Apple rootstocks. In: Rom, R.C. and Carlson, R.F. (eds) Rootstocks for
Fruit Crops. Wiley Interscience, New York, pp. 107–144.
Hatton, R.G. (1917) Paradise apple rootstocks. Journal of the Royal Horticultural Society 42, 361–399.
Lockhard, R.G. and Schneider, G.W. (1981) Stock and scion growth relationships and the dwarfing mech-
anisms of apple. Horticultural Reviews 3, 315–375.
Sadowski, A. and Hrotko, K. (1999) Apple Rootstocks for Intensive Orchards. Proceedings of an International
Seminar, Warsaw, 18–21 August 1999. Department of Pomology, Warsaw Agricultural University,
Poland, and Department of Pomology, University of Horticulture and Food Industry, Budapest,
Hungary, 131 pp.
Tubbs, F.R. (1972) Research fields in the interaction of rootstocks and scions in woody perennials, Parts 1
and 2. HortAbstracts 43, 247–253, 325–335.
Webster, A.D. (1995) Temperate fruit tree rootstock propagation. New Zealand Journal of Crop and
Wertheim, S.J. (1998) Apple rootstocks. In: Rootstock Guide: Apple, Pear, Cherry, European Plum. Fruit
Research Station, Wilhelminadorp, The Netherlands, pp. 19–59.
References
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6 Propagation and Nursery Tree Quality
S.J. Wertheim1 and Anthony D. Webster2

1FruitResearch Station, Randwijk, The Netherlands; 2Crop Science Department,
Horticulture Research International, East Malling, West Malling, Kent, UK
6.1 Introduction 126

6.2 Seed Propagation 126
6.3 Vegetative or Clonal Propagation 127
6.3.1 Stooling 128
6.3.2 Layering 129
6.3.3 Cuttings 131
6.4 Micropropagation 135
6.4.1 Propagation in vitro 135
6.4.2 Practical applicability 136
6.5 Tree Raising 136
6.5.1 Site choice 136
6.5.2 Planting distance 137
6.5.3 Rootstock cutting (heading) back and bleeding 137
6.5.4 Plant material 138
6.5.5 Tree support 139
6.5.6 Trunk cleaning 139
6.5.7 Other types of plant material 140
6.6 Budding and Grafting 140
6.6.1 Bud wood 141
6.6.2 Rootstocks, budding height and site 141
6.6.3 Budding and grafting methods 141
6.6.4 Tying and after care 142
6.6.5 Bench grafting 143
6.7 Branching 143
6.7.1 Manipulation of branching by hand 143
6.7.2 Manipulation of branching by chemicals 144
6.8 Defoliation and Digging Up (Lifting) Trees 146

126 S.J. Wertheim and A.D. Webster
6.1 Introduction Hereafter, the propagation methods for

apples are reviewed, with emphasis given to
In nature, apple trees multiply by seeds, but propagation on (moderately) dwarfing root-
when apple trees are grown commercially or stocks. For a successful modern orchard, the
by the home gardener the common propaga- planted tree must be of high quality, and
tion method is to bud or graft scion cultivars attention will also be paid to the raising of
on to selected rootstocks. The reason for trees capable of early cropping.
propagation by means of rootstocks is that Recent reviews on rootstock propagation
apple cultivars are not true to type when include Howard (1987) and Webster (1995).
propagated by seed and, at least in former
days, difficult, if not impossible, to propagate
by vegetative means. Thus, when a seedling 6.2 Seed Propagation
with good fruiting characteristics was found,
it could not be multiplied by sowing its seeds Although seedlings are still widely used as
or by use of layering or cutting techniques. rootstocks for raising apple trees, this use is
Today, vegetative propagation techniques on the decline in most countries. Trees
have improved so much that scion cultivars raised on the seedling rootstocks that are
can be multiplied by cuttings or by micro- currently available are too vigorous and
propagation in the laboratory. Nevertheless, variable for modern fruit growing.
the use of rootstocks has remained dominant, However, seedlings do have several advan-
because most of the currently grown scion tages compared with clonal rootstocks.
cultivars, once self-rooted, do not perform They are easily and cheaply produced,
well in the orchard. Scion trees propagated virus-free (as viruses are not transmitted
‘on their own roots’ are more vigorous, come through apple seed) and, if raised under
later into bearing and crop less efficiently certain conditions, free of soil-borne root
than those on good rootstocks. diseases, such as crown gall (Agrobacterium
Apple seedlings must have been the first tumefaciens (E.F. Smith & Townsend) Conn.).
rootstocks used extensively by early horticul- Orchard variability in the growth and crop-
turists, largely because seeds were readily ping of apple scions raised on seedling root-
available. Only after it was observed that stocks can be reduced by collecting seeds
some apple cultivars could be easily propa- from special seed orchards. Here, only a
gated by layering did vegetative or clonal tried and tested combination of two clonal
propagation come to be considered as a useful cultivars or one self-fertile cultivar is grown
means of propagating valuable scion culti- for seed production. ‘Delicious’ (USA),
vars. It must soon have become clear that ‘Antonovka’ (eastern Europe) and
‘clonal’ (vegetatively propagated) rootstocks ‘Bittenfelder’ (western Europe) are all rec-
had great advantages over seedling root- ognized seed sources, but other cultivars
stocks. First, there was the possibility of adapt- are used elsewhere. Seed plantations should
ing the performance of the scion cultivar in a be located far from other orchards to pre-
desired direction. This was important and vent unwanted cross-pollination. A further
remains especially so today for tree vigour, measure to reduce variability in perfor-
precocity and cropping efficiency. Secondly, mance of seedling rootstocks is seed grad-
orchards on clonal rootstocks were uniform in ing and the use of only the largest seeds, as
growth and cropping, while those on seedling these produce the best plants.
rootstocks were more variable because each Fruit from seed orchards is harvested by
scion tree is propagated on a distinct geno- hand or by mechanical shakers and the seeds
type. Today, there is a worldwide trend are separated from the fruit. Apple seeds
towards ‘intensive’ apple growing with scion cannot germinate directly after harvest and
trees grown on rootstocks that are more or extraction from the fruits, because they are
less dwarfing. Such intensive planting sys- dormant (Dennis, 1994). To remove seed dor-
tems are easy to manage and provide growers mancy, seeds must be stored for a certain
with early returns on invested capital. period at low, but above-freezing, tempera-
Propagation and Nursery Tree Quality 127
tures with adequate moisture and air – a The continuous use of ‘conventional’ (i.e.
process called stratification. The most effec- not apomictic) seedlings is partly due to
tive stratification temperatures for apple are their easy availability and is also attributable
2–6°C, with an optimum at 4°C. Ninety days to the rather conservative attitudes of certain
at 4°C can result in 100% seed germination rootstock suppliers and apple growers.
(Seeley and Damavandy, 1985). These figures Seedling-raised rootstocks of several
are indicative only and vary with seed Malus species do prove to be of value in
source, year and the fruit-storage regime improving the tolerance of apple trees to
(Perino and Côme, 1979). After stratification, drought conditions. Malus sieversii Ldb. or
apple seeds germinate best at temperatures Malus prunifolia Willd. seedlings are com-
from 10 to 20°C; at 30°C a secondary dor- monly used in parts of China with this objec-
mancy arises (Perino and Côme, 1977). tive and similar seedling rootstocks are used
Seed stratification is carried out in special in Japan. Where control of tree vigour is
cases in cold-storage rooms at 2–4°C. The required on seedling rootstocks, a dwarfing
seeds are placed either in sand or peat or interstock, such as M.26, is used. Seedling
without any substrate, but it is essential that rootstocks continue to be used, albeit much
the seeds do not dry out. After stratification, less than in the past. In the USA, they are
seeds are sown at about 2 cm depth. At an mainly used with compact scion cultivars of
early stage of growth the primary or tap ‘Delicious’, such as ‘Red Chief ’ or ‘Oregon
roots are mechanically removed by under- Spur’, where vigour control is achieved via
cutting the seedling rows. Planting distance the scion.
will depend on local conditions, the objec-
tives of the nurseryman and the machinery
available; a within-row distance of 20 cm is 6.3 Vegetative or Clonal Propagation
commonly used (Bärtels, 1982).
In the past, it was thought that apomictic New plants identical to a parent plant can be
seedlings of various Malus species closely produced by division (stooling or layering)
related to the cultivated apple might have a or by cutting techniques. In the former case,
future as rootstocks of very uniform perfor- young plants remain attached to the mother
mance. All apomictic seedlings are identical plant until they have formed roots and are
to the mother-parent cultivar, as they arise able to develop independently. Stooling and
from maternally derived cells within the layering are long-established methods of
ovary and not from the fertilized egg cell. division (Knight et al., 1928; Anon., 1963) and
Apart from giving uniform performance as are still the most common propagation meth-
rootstocks, apomicts are also virus-free. ods for apple rootstocks. With cutting tech-
However, the apple types used are only ‘fac- niques, young plant parts are separated from
ultative apomicts’. This means they produce the mother plant in summer or winter and
a mixture of apomictic and zygotic seeds are then induced to form roots. In the former
and the two must be separated in the nurs- case, leafy shoot tips collected in spring or
ery; this proves to be a cumbersome and early summer are rooted in glasshouses
often difficult activity. Most of the types of under high humidity; these are often referred
apomictic seedlings evaluated as rootstocks to as softwood cuttings. In the case of winter
also proved to be rather vigorous. Moreover, cuttings, leafless 1-year-old shoot parts are
when a scion cultivar contained latent collected and their rooting induced in a suit-
viruses and the apomictic rootstock type able medium with bottom heat. Occasionally,
was hypersensitive, incompatibility could semi-hardwood cuttings, taken in late sum-
occur, although this defect can be solved by mer once shoot growth has terminated, have
using virus-free scion material (Schmidt, been used for propagation of apple root-
1988). For these shortcomings and because stocks, although this is not a common prac-
of the strong market demand for dwarfing tice. Micropropagation is another, relatively
clonal rootstocks, apomictic seedlings are new, method used for propagating both root-
not used in commercial apple growing. stocks and scions. Here, tiny plant parts are
separated from the parent plant, for example takes place. The number of times that shoots
growing shoot tips, and multiplied and are earthed up varies and depends on local
rooted on special media (in vitro) under ster- practice. As well as soil, peat mixtures or
ile conditions in the laboratory. All the above sawdust are also used for earthing up in the
methods have their strong and weak points stool beds. However, care must be taken to
and all can be used for the propagation of ensure that the sawdust used contains no
both scion and rootstock cultivars. Given the substances inhibitory to rooting.
shortcomings of self-rooted scion cultivars, After natural leaf drop, the ridge of soil
the methods are mainly used in rootstock or other substrate is ploughed and/or
propagation. Whatever the method of propa- forked away and the rooted shoots are cut
gation, it is recommended that, where avail- loose from the parent plants. Depending on
able, only virus-indexed material that is the climate, the removed soil or substrate is
healthy and true to type should be used. To raked back over the stools after harvest to
ensure healthy and true-to-type stock, the protect them from winter injury. The above
safest course of action is to use only material cycle of production is repeated annually
obtained from organizations that have been and, provided that no diseases or pests
officially commissioned to maintain and dis- interfere, the stock plants should continue to
tribute healthy propagation materials. produce rooted shoots of adequate quality
for at least 15 years.
Rootstock clones differ in the number of
6.3.1 Stooling rooted shoots (‘liners’) that are produced
annually per stool plant or per metre stool
With stooling (stool or mound layering), 1- bed. Average annual yields of first- and sec-
year-old rooted plants are planted vertically in ond-grade shoots obtained at East Malling
spring and left unpruned for 1 year. In the fol- Research Station varied between 2.6 per stool
lowing spring, the stems are cut back to 2–3 for M.9 to 13.4 for MM.104 (Howard, 1977).
cm above the ground and the arising shoots Production figures from trials conducted ear-
are partly covered with earth several times lier on 4-year-old stools planted at 60 cm
during the growing season. This is mainly apart in the row at the same research station
done mechanically but, if needed, additional ranged from a total of nine plants (of which
handwork can be involved, especially at the four were first-grade liners) per stool for M.9
first covering (‘earthing up’). Covering the to a total of 24 (with 18 first-grade liners) for
shoot bases must be done carefully and in M.13 (Knight et al., 1928).
such a way as to leave a sufficient amount of The productivity of stools is negatively
the leaves exposed to the light but at the same affected by the presence of viruses
time to blanch the basal part of the stem to (Campbell, 1961) or by using liners that are
facilitate rooting. Severe pruning and blanch- too small to establish the stool beds, even
ing are both essential for successful rooting in when a 2-year establishment period is used
the stool bed (Howard et al., 1985). (Howard, 1977). By biennial harvesting, the
To allow earthing up, distances between number of large liners produced, which are
rows of stools should be adequate – at least 1 needed where high budding is to be prac-
m. Within the rows, stools are spaced 30 cm tised, can be increased. This method
apart. When the young shoots are about decreases the number of non-rooted shoots,
10–15 cm long, friable soil is carefully drawn welcome in the case of shy-rooting root-
up to the rows and in between the plants, so stocks, such as M.27. However, negative fea-
as to cover up shoots to half their total tures of biennial harvesting are the increase
lengths. A second earthing up is done when in plants with lateral shoots, which are a nui-
the shoots are approximately 20–25 cm long sance when budding, and the number of
and a final one when shoots have grown to misshapen and mildew-infected (Podosphaera
about 45 cm. After this final cultivation, leucotricha (Ell. & Evi) E.S. Salmon) plants
15–20 cm of the shoot bases should be cov- (Vasek and Howard, 1984). A way to
ered with soil and it is there that the rooting improve the performance of stool beds is to
kill the weak growths that develop early in the horizontal layers by the circular saws
the season on stumps of weak shoots left used at harvest, the land surface must be
after harvesting and pruning in the previous completely flat with all plants maintained in
year. At about 5 cm length, these shoots can a horizontal plane.
be easily eliminated using one spray of 4% In The Netherlands, the procedure used
‘Tipp-off ’ (naphthaleneacetic acid (NAA) for layering M.9 is as follows. The plants are
plus decanol and emulsifiers). Provided the planted obliquely at 30 cm spacings in the
growth of the stool plants is vigorous, more rows, which are 75–100 cm apart (Plate 6.1).
large-rooted shoots suitable for high bud- November is the optimum planting time, but
ding can then be harvested (Howard, 1984). this depends on weather and soil conditions.
The same objective can also be achieved by For establishment, plants with a stem diame-
removing the weak shoots by hand. ter of 8–10 mm measured at ground level are
The stool harvest consists of ‘liners’, ‘non- used. After one growing season, the plants
liners’ and non-rooted plants. Liners are are forced horizontally by braiding (tying)
rooted plants of adequate size suitable for them together, including any side-shoots that
lining out in the fruit-tree nursery for bud- may have arisen. Given the close planting
ding. Non-liners are rooted but too thin for distance in the rows, there is an overlap of
budding in the following spring/summer. shoots from adjacent layers that facilitates
Both, non-liners and the non-rooted plants braiding and contributes to high production.
that are not too small are planted (‘bedded The first earthing up is carried out when the
in’) at close between-plant spacings (7.5–10 shoots are 20–25 cm long and either soil or
cm) for a further year for additional thicken- peat mixtures are used. The peat is brought
ing and/or rooting. They may be used for into the rows using machines, but handwork
late-summer budding in these ‘waiting’ beds may be needed for distributing soil or peat
or lined out in the subsequent season. Very evenly between the shoots. At intervals of
small rootless plants are usually discarded. approximately 1 month, two further earth-
Apple-scion cultivars can also be success- ings up are undertaken. On these occasions,
fully multiplied by stooling (Schimmelpfeng, soil is usually used and the shoots are par-
1963), although this is rarely practised com- tially covered to leave half of their lengths
mercially. exposed. The rooted shoots are cut off with a
circular saw before winter (Plate 6.2) but
after leaf drop, and are stored indoors at 1°C
6.3.2 Layering at high humidity. The saw cuts the shoots at
about 1 cm above the horizontal layers and
One-year-old plants are planted in rows, the below-ground blanched section should
which are preferably north–south-orientated, be at least 15 cm long. When properly fertil-
at an oblique angle of 30–40° to the horizon- ized and managed, especially in regard to
tal and left unpruned for one growing sea- crop-protection sprays, layer beds continue
son. The following spring, the stems are bent to produce well for more than 15 years.
and secured flat to the ground and the In practice, 20–25 well-rooted plants of
shoots that arise on these horizontal layers more than 5–6 mm basal stem diameter per
are regularly and carefully earthed up, just metre of row length can be obtained, plus
as with stooling. In trench layering, the 15–20 plants that are too thin for immediate
mother plants are planted in a shallow use as liners (less than 5–6 mm diameter).
trench so that when bent horizontally the These thin rootstocks are planted for another
layers are just below ground level. In this year at close spacings (10 cm in rows 50 cm
method, which is often used with stone apart) in a waiting bed. M.9 plants destined
fruits, the layered shoots can be covered with for budding are of various basal stem diame-
a few centimetres of earth just before bud ter classes (5–7 mm and 7–9 mm or 6–8 mm
break. For apple, however, this is not neces- and 8–10 mm), depending on the particular
sary and the mother plants are usually practices of the nursery. Plants over 9 or 10
planted on flat land. To prevent damage to mm in diameter are used for bench grafting.
Yields of layer beds depend on the grow- gation method also had an effect; in vitro
ing conditions (soil and climate), age, health plants were more productive than those
(especially virus) status, rootstock clone (or obtained by conventional layering (see sec-
even sub-clone) and efficiency of manage- tion on micropropagation). It should be men-
ment. Layer beds start to produce in the sec- tioned that both the in vitro and the layer
ond season following planting, and plants of each of the three sub-clones origi-
production increases in the following few nated from a single plant. Material of that
years before stabilizing at a constant output plant was either put through in vitro culture
(Plates 6.3 and 6.4). The difference in the pro- and thereafter the shoots produced used for
duction level of a layer bed at an early and in layering or the plants were solely propa-
a mature stage is approximately 30% (Table gated via a layer bed. These differences in
6.1). The 8-year average gives an indication origin did not affect subsequent orchard per-
of the performance of M.9 layers of different formance (Wertheim and Kunneman, 1993).
origins. Juvenile sub-clones of this rootstock, With P.22, similarly, in vitro culture pro-
such as Fl.56 and RN.29, are more productive duced juvenile plants clearly different from
than the non-juvenile T.337. A virus-infected those obtained solely through layering.
sub-clone (B.984) gives poorer production. Compared with the adult type, the juvenile
Approximately 40–50% of the yield consists sub-clone was characterized by smaller
of plants suitable for budding and about 5% leaves, more laterals, a spreading habit, a
for bench grafting (Table 6.1). The rest must higher production per metre layer bed and a
be grown on for another year. higher rooting percentage. However, the
The occurrence of M.9 sub-clones in juve- pendulous habit of the juvenile sub-clone
nile, adult and transitional stages is due to was a nuisance for nursery management
their maintenance in certain ontogenetic (Wertheim, 1991). The differences in sub-
stages (van Oosten, 1986). Production of clones had no consequences for the subse-
M.26 has also been shown to vary with the quent orchard behaviour of trees raised
sub-clone; juvenile types again producing upon them. A good sub-clone for the root-
more than adult ones (Table 6.2). The propa- stock propagator, however, need not neces-
Table 6.1. Production of a layer bed (in plants per metre row length and greater than
4 mm diameter) of virus-free M.9 sub-clones planted in spring 1983 at 1.25 × 0.25 m
between- and within-row spacing. (Data from Versteegen and Verstraelen, 1992.)
Sub-clones
Parameter T.337 Fl.56a RN.29a B.984b
Average 1984/85 20.5 22.3 22.7 16.9

Average 1990/91 31.6 34.6 42.8 25.5
Average 1984–1991 26.6 29.0 32.5 21.3
Averaged for 1984–1991
Rooted plants (%) 89 94 94 86
Spurred plants (%) 6 7 13 3
Plants 4–6 mmc (%) 46 50 44 47
Plants 6–8 mmd (%) 33 33 33 33
Plants 8–10 mmd (%) 15 11 16 14
Plants 10–12 mme (%) 5 3 5 5
Plants 12 mme (%) 2 2 2 1
aJuvenile sub-clone.
bNon-virus-free.
cFor bedding.
dFor budding.
eFor bench grafting.
NB. Spurred plants have short laterals.

Table 6.2. Production of layer beds (in plants per metre row length) of virus-free M.26 sub-
clones planted in spring 1989. Plant material of each sub-clone was propagated by layering,
but the layer beds were established either from a maternal layer bed or from in vitro culture,
using one single plant as the source of all plants. (Data from Denissen et al., 1993.)
Propagation Plants m−1 Stem diameter (mm)

method Sub-clone Character 1991 1992 1991 1992
Layering only IVT Non-spurred 16.0 25.0 9.2 7.3

NAKB Non-spurred 19.4 27.0 8.5 7.1
NAKB Spurreda 26.1 37.4 8.8 7.0
Average 20.5 29.8 8.8 7.1
Layering IVT Non-spurred 18.5 26.2 9.4 7.7
after in vitro NAKB Non-spurred 24.5 35.2 8.7 7.0
NAKB Spurreda 33.4 49.0 8.2 6.7
Average 25.5 36.8 8.7 7.0
aJuvenile type.
sarily be the best one for the following users more difficult-to-root rootstocks, such as M.9
of that material (including the raisers of and M.26 (Doud and Carlson, 1977).
scion trees and fruit growers). Juvenile Shoots derived from stools or from layers
clones may have more spines or laterals, are not completely uniform. This may be
which are a nuisance with budding, and because of their different position of origin
may be more prone to sucker and burr-knot relative to the roots. Those arising close to
formation in the orchard. Suckers can be the roots may be slightly more vigorous and
points of entry for pests and diseases; burr bear more laterals or spines than those origi-
knots have reduced cold-hardiness and nating more distally. Root hormones may be
cause tree-to-tree variability. involved here, although there is little evi-
Compared with M.9, layer beds of other dence to support this hypothesis. In a Dutch
popular rootstocks, such as M.26, MM.106 trial, orchard performance of M.27 rootstocks
and MM.111, can be more productive, those separately harvested from basal or apical
of M.4, M.7, M.27, J.9, B.490, B.491, P.16 and parts of layers was similar. The former were
P.18 as productive and those of P.1, P.2, P.22, slightly more vigorous and had a few later-
B.9, O.3 and M.25 less productive (Quamme als, while the latter had none.
and Brownlee, 1990). It should be realized,
however, that yields may be affected by the
sub-clone chosen. 6.3.3 Cuttings
In both stooling and layering, the emer-
gence of roots appears to be largely confined Propagation of apple rootstocks or scions by
to nodal positions near lateral buds, proba- hardwood or softwood cuttings is not wide-
bly because the mechanical restriction there spread. Stooling and layering techniques are
is the least. Blanching (i.e. partial etiolation) currently perceived to be more reliable and
causes an increase in starch in the parenchy- less expensive. Details on cutting methods
matous bud and leaf gaps and in the outer are given by Hartmann et al. (1990) and on
ring of pith cells of the stem, together with a stock-plant manipulation by Howard (1994).
decrease in the degree of sclerification of the
cortex. A high starch content favours root
6.3.3.1 Hardwood cuttings
formation, perhaps because it provides more
energy, and rooting appears to be negatively Propagation by hardwood cuttings is some-
correlated with the degree of sclerification. times seen as a preferred alternative to lay-
Easy-to-root rootstocks, such as MM.106, ering, because shoots can be harvested
have a lower percentage of sclerification than from stock plants each year in numbers
determined by demand. However, hard- preferred method of application is to dip the

wood cuttings of apple rootstocks usually cutting bases for 5 s into a 1 cm deep solu-
require growing in the nursery for a year tion of 2500 p.p.m. IBA in a 50% aqueous
before they can be budded (Plate 6.5). This solution of acetone, but other methods and
means that demand for rooted liners suit- carriers can be used. Too high concentrations
able for budding must be known 2 years in of auxin may contribute to basal rotting and
advance if the technique is to have signifi- should be avoided. After the auxin treat-
cant advantages. Research, mainly carried ment, cutting bases dry naturally before
out in England, has shown that it is possi- being inserted into a rooting medium. Auxin
ble to root hardwood cuttings of apples, treatment also stimulates successful estab-
provided that the bases are treated with lishment of the cutting. Optimal establish-
auxin and given a brief treatment at high ment is usually achieved by planting in
temperature. Wounding the base of the cut- February (northern hemisphere). Visible
ting also has favourable effects on rooting roots are not necessary on transfer to the
(Howard, 1987). field, but the presence of healthy callus is
Current-year shoots are cut after leaf fall (Howard, 1968, 1978, 1985, 1987; Howard et
in autumn or early winter from hard-pruned al., 1984; Campen et al., 1990).
stock plants. These should be virus-free, as In the rooting medium, the cutting bases
the presence of viruses reduces shoot pro- are subjected to a period of relatively high
duction, subsequent root formation and temperature. The heat treatment operates
establishment (Howard, 1972). The stock additively to and independently of the auxin
plants are planted in hedgerows at distances treatment. A suitable medium is needed to
suitable for management by tractors or other minimize rotting, and mixtures of peat with
machinery. Distance between stock plants in adequate amounts of coarse-particle materi-
the rows should be at least 30 cm. After als, such as sand, grit or perlite, are satisfac-
removal of the cuttings from hedge plants, tory. The medium should be able to hold the
the remaining shoots are shortened to three cuttings in place, provide moisture, permit
or four buds to ensure vigorous growth in air penetration and gas exchange and create
the subsequent year. Shoots from vigorous a dark environment suitable for rooting.
plants have a higher rooting potential than Good drainage of the rooting medium is a
those from lightly or non-pruned plants and necessity because, although hardwood cut-
those from low hedge plants root better than tings take up water from the medium, they
those from high plants. Thus, hedges are cannot transpire it rapidly as they have no
kept at about 1 m in height (Plate 6.6). The leaves. As a consequence the cutting bases
rooting potential is highest at the shoot base can rot if inserted into medium that is too
and this part must be included in the cutting. wet and/or poorly drained. At Horticulture
When the cutting includes the basal ring of Research International (HRI), East Malling,
buds near the old wood it is called ‘basal an 8 cm layer of granulated pine bark over
nodal’, when the cut is made a node higher 20 cm fine sand is successfully being used
‘non-basal nodal’ and if cut between this and with a heating mat situated on top of the
the following node ‘non-basal internodal’ sand. Bundles of cuttings are put into the
(Howard et al., 1984). Cutting length bark so that their bases are about 3 cm above
depends on planned planting depth and the heating elements. The optimum tempera-
budding height and varies between 40 and ture of the rooting medium at the cutting
60 cm. Cuttings are shortened distally to the base depends on the cultivar, but in general
required length. Growing cutting hedges in a 20°C is recommended for apple (Howard,
polyethylene tunnel increases vigour, rooting 1987). However, the apple scion cultivar
percentages and roots per cutting (Campen ‘Cox’s Orange Pippin’ reacts better to 30°C
et al., 1990). (Webster et al., 1990). The duration of the
Applying auxin to the cut surface greatly heating period depends on the time of the
enhances rooting. The most effective auxin is year and the cultivar. In February, 2–3 weeks
indole-3-butyric acid (IBA). For apple, the suffice, while in midwinter 4 weeks are bet-
ter. Easy-to-root cultivars need less time such primordia are present, but not in M.26
than difficult-to-root ones. For the former, a and M.9. In MM.106 and Mo84, primordia
good indication of successful stimulation is are present in the cortex around the bud and
the presence of healthy callus and only a new roots emerge from that area. The pres-
small percentage should show a few roots. ence of preformed roots is an advantage in
Difficult-to-root cultivars require the pres- propagation by cuttings. In M.26 and M.9,
ence of a few roots on all cuttings for good roots have to be formed from newly formed
subsequent establishment The rooms where callus at the basal cut surface. In Mo84, roots
cuttings are treated (in what are often called can emerge both from callus and from buds
‘Garner bins’) have minimum light and 90% and lenticels, which is favourable for hard-
humidity and are cool (max. 10°C) to pre- wood-cutting propagation in the field.
vent bud break. Under humid conditions, When the basal cut surface rots, roots can
cuttings root best in late autumn and again still emerge from lenticels and buds above
in late winter and spring, but less well in the rotted area. In the new dwarfing root-
midwinter. Under 50–60% humidity this stock Q9, root primordia occur in both nodal
trend is reversed. After heat treatment, cut- and internodal areas and this will further
tings can either be planted in the nursery for contribute to easy propagation (Fukuda et
budding in the summer or in waiting beds al., 1988). Differences in hormone metabo-
of compost to establish further (Howard, lism may further contribute to the variable
1987; Webster et al., 1990). rooting ability of apple cultivars (Hartmann
Rooting is stimulated by wounding, but et al., 1990).
only in the presence of auxin. Splitting the Hardwood cuttings are being used to
shoot base for a length of 2 cm is especially some extent for easy-to-root apple root-
effective (Howard et al., 1984; Howard, 1987, stocks, such as M.26, M.27, MM.106 and
1994). Possibly, this splitting acts by breaking MM.111, and for non-rooted shoots from
the sheath of sclerenchyma tissue that blocks stool or layer beds. In future, more root-
root emergence. In apple shoots, such a stocks will certainly be added to this list.
sheath of fibres and sclereids occurs in the
primary phloem. Difficult-to-root apple culti-
6.3.3.2 Softwood cuttings
vars have a more complete sheath than eas-
ily rooting ones. The development of Softwood cuttings are quite small in size, to
sclerenchyma tissue takes place during shoot minimize water loss, and are therefore less
growth. It may vary between apple cultivars suited to rootstock production. This is
and is subject to variation in the environ- because it takes too long before rootstocks
ment. Formation is retarded by shading and suitable for sale or for budding are obtained.
etiolation. Partial etiolation occurs in earth- Softwood cuttings can, however, be useful
ing up of stools and layers and in banding for the rapid multiplication and build-up of
shoots with opaque tape (blanching), and a new rootstock cultivar (Howard, 1987).
these measures enhance rooting. If the devel- Softwood or summer cuttings are also of
opment of the sheath is rapid and is near the practical value in certain specific situations.
stem tip, rooting of summer cuttings is diffi- In Norway, import of rootstocks is prohib-
cult. Although it takes longer, rooting of ited for phytosanitary reasons and the grow-
young shoot tips under mist is possible ing season is too short for successful layering
(Beakbane, 1961, 1969; Pontikis et al., 1979) (Billing Hansen, 1990).
(see paragraph on summer cuttings). Softwood cuttings are young extension
However, other factors than the scle- shoots that are cut in spring or early summer
renchyma sheath are involved in differences from hard-pruned stock plants. Shoots from
in rooting potential among cultivars. For stools root better than those from hard-
example, the presence of preformed root pri- pruned 1 m high hedges. It appears that the
mordia in the cortex varies between culti- rooting potential of the cuttings increases
vars. In M.7, M.27, MM.106, Mo84 and the with increasing severity of pruning and
new dwarfing rootstocks Q9, Q60 and Q64 decreasing distance between the position of
the cutting on the stock plant and its root Hansen, 1990), but the solution-dipping
system (Nelson, 1976; Howard et al., 1985). method generally seems better. It is recom-
Stock plants derived from in vitro material mended that cuttings be treated with a
produce cuttings that root more easily than fungicide before ‘sticking’ them into a root-
conventionally propagated stock plants ing medium – often a mixture of peat, sand
(Quamme and Hogue, 1994). Also, stock and grit. Wounding of the cutting bases does
plants that have been kept in the shade for a not stimulate rooting.
while from bud break onwards give better The loss of water from the cuttings should
results than plants growing in full daylight. be minimal. This can be achieved using
The first cuttings obtained from stock plants intermittent mist (IM) or fogging systems.
in the growing season give higher rooting The driving force resulting in water loss by
percentages and numbers of roots per rooted the cutting is the difference in pressure
cutting than later collections (Delargy and between water vapour in the leaves (Vleaf )
Wright, 1978; Howard et al., 1985; Billing and that in the surrounding air (Vair). This
Hansen, 1989, 1990; Grzyb et al., 1989). difference should be kept as small as possi-
Cuttings are preferably made early in the ble for success in rooting. IM mainly acts by
morning when shoots and leaves are turgid. decreasing Vleaf, through reduction of leaf
To prevent wilting, transfer of the cuttings to temperature, and partly by a modest
the rooting environment (glasshouse or poly- increase in Vair. IM is only applied in the
ethylene tunnel) must proceed quickly and, daytime. Under sunny conditions, every 5
if necessary, the cuttings must be dipped into min water is supplied for 3–4 s. With over-
water and transported in plastic bags con- cast weather, mist is applied less frequently,
taining some water. for example, every 10–30 min, but other time
Various shoot parts and various lengths regimes are possible. A fog system maxi-
of cutting have been successfully used with mizes Vair by raising the ambient humidity.
apple, as illustrated by the following exam- In apple, a fog system gives better results
ples. Shoots can be collected in late June than IM. Fog is supplied when relative
(northern hemisphere) from outdoor stock humidity falls below 90% and needs auto-
plants using the proximal 20 cm with three matic continuous control (Billing Hansen,
to four leaves. These are trimmed to a node 1989; Hartmann et al., 1990).
and deleafed at the lower end for about 7 cm Bottom heat (20°C) is not strictly neces-
(Howard et al., 1985). Short shoots up to 8 cm sary for the rooting of softwood cuttings, but
long developing from forced stock plants in may improve rooting (Nelson, 1976).
a glasshouse cut at their base can also serve Suitable air temperatures in the rooting envi-
as good starting material (Billing Hansen, ronment range from 20 to 25°C. The rooting
1990), as can regrowths just exceeding 13 cm process takes from several weeks to several
from rootstocks pruned at ground level months. When the cuttings are rooted, they
(Nelson, 1976). In all these cases, the lower should be hardened off by gradually dimin-
leaves are removed. Four-node cuttings per- ishing the air moisture content. Cuttings
form better than smaller ones, possibly deteriorate when left under mist too long
because their reserves are greater after they have rooted. The subsequent
(Schmadlak, 1969). In the case of long cut- growth of the cuttings depends on the time
tings, the proximal halves root better than of the season when rooting takes place, i.e.
distal ones (Quamme and Hogue, 1994). the time left for growth after rooting.
Exogenous auxin is necessary for the root- Climatic conditions and planting distance
ing of softwood cuttings and more so for dif- also play a role. After forcing stock plants
ficult-to-root cultivars than for easy-to-root from the end of March, planting rooted cut-
ones. The cutting bases are dipped for 5 s in tings in May (northern hemisphere) at 50–70
a 50% aqueous acetone solution containing plants m−2 gave the highest percentages of
2500 p.p.m. IBA to a depth of 8 mm and saleable rootstocks (diameter 6 mm) – 65%
allowed to dry (Howard et al., 1985). One per in the case of MM.106 and 45% for M.26
cent IBA in talc is also successful (Billing (Billing Hansen, 1990).
Many apple rootstocks have been propa- 6.4 Micropropagation

gated through softwood cuttings, but the
degree of success depends on the cultivar. 6.4.1 Propagation in vitro
For instance, MM.106 responds better than
M.26 (Billing Hansen, 1989) and M.7 better Apple rootstock and scion cultivars can also
than M.4, M.9, MM.104, MM.106, MM.109 be propagated in the laboratory. The most
and MM.111 (Schmadlak, 1969), B.9 is easier common method is to use tips or parts of
than P.2 or P.22 (Grzyb et al., 1989) and, shoots of a few mm to 1 cm in length. These
within the Ottawa series, O.7 is the most tiny ‘explants’ are surface-sterilized and cul-
difficult one (Nelson, 1976). Without forcing tured in vessels on a suitable medium in illu-
stock plants, softwood cuttings are not minated rooms under sterile conditions. The
likely to have much future for commercial various media used contain mineral salts, a
rootstock propagation, given the long time carbohydrate source (sucrose or sorbitol) and
period needed to produce saleable root- a cytokinin, often 6-benzyladenine (BA). The
stocks. shoots, once established in vitro, grow and
Difficult-to-root apple scion cultivars, produce new shoots from axillary buds (Plate
such as ‘Bramley’s Seedling’, can also be 6.7). Shoots are excised at intervals of about a
propagated by softwood cuttings, but only month and placed into a fresh medium for
with a combination of three techniques, further multiplication. This subculturing can
namely, temporarily covering the stock go on for years. When sufficient shoots have
plants with black polyethylene and then proliferated, the small shoots are induced to
form adventitious roots (Plate 6.8) on
taping and bark-ringing the shoot bases.
another, cytokinin-free, medium, which
Covering of the plants is done from bud
should contain an auxin, usually IBA (Jones,
break until the shoots are 8–10 cm long.
1993). There are several variations and modi-
Thereafter, 7.5 cm of the shoot base is taped
fications in the media and in the procedures
with a band of black polyethylene to keep
successfully used for apple culture in vitro
this part of the cutting etiolated. When,
(Lê, 1985; Collet and Lê, 1987, 1988;
after a few weeks, shoot length reaches
Hutchinson and Zimmerman, 1987; Pawlicki
20–30 cm and the etiolation effects of the
and Welander, 1994). Rooting can also be
covering have disappeared and the leaves
stimulated by adding certain thiol com-
are green again, the tape is removed and
pounds to the medium (Auderset et al., 1996).
the bark is ringed (4 mm) proximal to the When sufficient roots have been formed, the
basal blanched stem part. The combination plants are transferred to pots filled with pot-
of all three treatments can lead to high root- ting compost or another substrate and
ing percentages and good numbers of roots brought into a glasshouse for further devel-
per cutting (Delargy and Wright, 1978). opment. Establishment in the glasshouse also
With rootstocks, simply banding the shoot depends on the potting substrate (Lê and
bases with Velcro (2.5 cm × 2.5 cm) for up Collet, 1991) and can be improved by using a
to 20 days before the cuttings are collected fog installation and elevated carbon dioxide
can also improve rooting percentages and levels (van Telgen et al., 1992). Finally, the
root numbers per rooted cutting – for young plants are planted in soil outdoors.
example, with M.9 and MM.106 rootstocks. Because this propagation method is quite
The establishment and subsequent growth complicated and costly, quicker methods
of M.9 are improved and rotting of the have been sought. An important simplifica-
stem bases is diminished by this Velcro tion is to cut out rooting under sterile condi-
band (Sun and Bassuk, 1991). Blanching of tions and to root the shoots directly into the
non-earthed-up stool shoots with opaque potting substrate after dipping their bases in
tape is more effective than earthing up, IBA powder (Simmonds, 1983) or after
probably because it is done earlier and another short auxin treatment in vitro
excludes the light more effectively (Howard (Auderset et al., 1994). This method, some-
et al., 1985). times referred to as ‘direct sticking’, also
improves establishment in the soil, often a propagated plants, but without the disad-
weak point after in vitro propagation vantages of rejuvenation (Navatel et al., 1988;
(Webster and Jones, 1991). The establishment Webster and Jones, 1992; Jones and Webster,
problems are possibly due to the modified 1993; Czynczyc et al., 1994; Grant and
root anatomy of roots formed in vitro Hammatt, 1999; see also Table 6.2).
(McCleland et al., 1990), although the stom- A short supply of suitable rootstocks and
atal closure of leaves formed on in vitro- the long duration involved with tree raising
raised shoots is also often impaired and this have also been stimuli for in vitro rootstock
can result in their rapid desiccation. propagation and even tree raising. Using
The production of shoots and roots on in such techniques, it is possible to raise a
vitro cultures usually increases with ongoing branched tree on a rootstock in as little as 1
subculturing. The gradual physiological year (Hogue and Neilsen, 1991). However,
change during subculturing leading to these this method requires more skill and equip-
improvements is termed ‘rejuvenation’. It ment than traditional methods and has not
was assumed that the number of subcultures been followed up in practice.
was important (Webster and Jones, 1989) in If in the future scion cultivars become
this rejuvenation, but later it was shown that available that are compact and yield-produc-
the total time spent in culture was the deci- tive on their own roots, in vitro propagation
sive influence (Grant and Hammatt, 1999). may replace propagation via rootstocks.
Cultivars, but also clones of one cultivar, dif- Preferably, compact cultivars should not have
fer in their suitability for micropropagation, a chimeral stucture, as is currently evident in
and within a cultivar differences even occur some existing spur types, because in vitro
between shoot-culture lines originating from propagation may result in reversions back to
different shoot-tip explants (Webster and the original more vigorous parent type, so
Jones, 1989, 1991; Collet et al., 1994). leading to variable orchards. It has been sug-
gested that reversion may be overcome by
reculturing the compact type (Zimmerman,
6.4.2 Practical applicability 1997). Although, so far, micropropagation has
made no great impact on commercial propa-
To date, micropropagation plays only a very gation practices for apple, propagation in vitro
minor role in practical apple propagation. through adventitious regeneration remains of
For scion cultivars, this is due to their poor the utmost importance for the understanding
performance in the orchard (Buban et al., of fundamental physiological processes and
1993), in particular their delayed precocity of for aiding new methods of crop improvement
cropping (Plate 6.9). Part of this delayed via biotechnology (Jones, 1993; Diekmann et
cropping of trees derived from in vitro cul- al., 1999; Zhu and Welander, 1999).
ture may be due to their small size at plant-
ing compared with conventionally
propagated trees (Webster et al., 1985). Also, 6.5 Tree Raising
self-rooted trees derived from micropropaga-
tion can suffer badly from root suckering 6.5.1 Site choice
and burr-knotting. This may also occur when
rootstocks originating from micropropaga- In order to achieve the level of growth suffi-
tion are used directly as liners for budding cient to raise high-quality trees in the nursery,
with scions. Therefore, the only current prac- it is essential that the soil is of high quality
tical use of micropropagated material is for and well drained and that the site has a provi-
establishing stock plants of rootstock clones, sion of irrigation. It is also of the utmost
i.e. as starting material for cutting hedges, importance that the soil has never before been
stool beds or layers. For many years, such used for apple cultivation. On ‘fresh’ (virgin)
material produces many more cuttings or soil the growth level of young trees is much
liners per plant with improved rooting better than where trees have been grown before
capacity compared with conventionally (‘replant sites’). In many areas nurserymen
are still allowed to treat the soil with chemi- good tree development. To facilitate good
cals against these ‘replant problems’ and such light interception by the leaves and to ensure
treatments do improve young tree growth. an even uptake of water and nutrients by the
On light soils, the nematode Pratylenchus pene- roots, a square planting system seems better
trans (Cobb) Filipjev & Schum.-Stek. is mainly than an extreme rectangular one. Indeed, ade-
responsible for poor and variable growth and quate between-tree distance within the row is
various nematocides are used to control it. On important for side-shoot formation (‘feather-
heavy soils, other pathogens are involved in a ing’) and for good tree-stem diameter (Table
‘specific replant disease’ and other fumigants 6.3). For trees on M.9, a planting distance of
are needed. However, increasingly approval 100 cm × 35 cm is ideal on most fertile soils
for the use of soil fumigants is being with- and will produce trees of good quality and at
drawn as part of tightening pesticide legisla- reasonable tree numbers per hectare. For
tion. Therefore, it is better to avoid replant more vigorous rootstocks, increased tree spac-
problems altogether and to secure fresh land ings may be needed, but no data are available
for every nursery cycle. Even after fumiga- to support this suggestion.
tion, tree growth on soils previously cropped
with apples rarely matches that achieved on
virgin soils. 6.5.3 Rootstock cutting (heading) back and
bleeding
6.5.2 Planting distance Established budded rootstocks are cut back

above the bud grafts in the spring following
Whatever the method of tree raising that is budding. When this is done in winter, trees
employed, an adequate distance between the may develop better than when cutting is
rootstocks in the nursery is needed to allow done in spring. However, in certain years
Table 6.3. Tree-quality parameters of ‘Jonagold’ (J) and ‘Gloster’ (G)a on M.9 rootstock at various
planting distances in the nursery. Rootstocks (6–8 mm diameter) planted spring 1985 at 16 different
spacings (combinations of four row and four tree spacings). Budding August 1985. Measurements
autumn 1986. (Adapted from Wertheim, 1986a.)
Stem diameter Laterals per tree

Tree height 10 cm above higher than 40 cm Total lateral length
Spacings (cm) above union (cm) union (mm) above soil level (no.) per tree (cm)
Within-row tree spacings: data averaged for row distances

15 121.7 9.4 2.0 41.5
30 124.7 11.0 4.7 147.8
45 125.0 11.2 5.7 202.0
60 125.8 11.9 5.8 207.8
F test NS * * *
LSD0.05 – 0.7 1.5 72.5
Between-row spacings: data averaged for tree distances

60 121.9 10.5 4.3 129.5
70 124.1 10.9 5.0 157.2
80 127.7 11.4 4.4 146.9
100 123.0 10.8 4.9 165.8
F test NS NS NS NS
LSD0.05 – – – –
aNo laterals with ‘Gloster’.
F test: cultivar × distance NS; tree distance × row distance NS; thus 16 tree spacings combined.
NS, not significant; *, significant (P 0.05).
and on light soils winter pruning can cause bud union is available for sap loss. Later, the
severe tree losses due to the ‘bleeding syn- stumps left by both methods are removed
drome’. Rootstocks may die back or buds directly above the bud. A temporary snag
may fail to break or, if they do, the shoots has no negative effects on tree quality. Soil
may wilt later on, and the rootstock bark cultivation to a depth of about 10 cm,
becomes spongy or papery. It is assumed destroying part of the surface root system, is
that, when the tissues are still dormant, cut- another suggested remedy. If bacterial infec-
ting back dehardens the plant. This is dan- tion of the cut surface occurs, sprays with
gerous as in spring many frosts may still bactericides might alleviate the problem, as
occur and at ground level temperatures may will providing a temporary straw cover after
drop to very low levels. These freezes may cutting back as protection against frosts.
easily damage dehardened tissues. The most radical solution is to completely
Moreover, by pruning, the rootstock stem abandon tree raising with established root-
has become very small relative to the root stocks on light soils. Indeed, with snip or
system. As the sun warms up the soil, roots interstem tree raising, which starts with
become active and take up water, root pres- bare-root rootstocks, fewer bleeding prob-
sure is built up and the cut rootstock may lems are experienced.
start bleeding from the pruning wound.
When the wound is dried out after early cut-
ting, the sap escapes through the rootstock 6.5.4 Plant material
bark. On heavy soil, bleeding is normally not
excessive. Compared with light soil, water The majority of apple trees produced in nurs-
uptake requires more energy and soil tem- eries are of one of three types: 1-year-old
peratures are not so readily raised. On light (‘maiden’) trees, ‘snip’ trees and interstem
soils, therefore, heavier tree losses occur due trees. One-year-old trees originate from bud-
to early cutting back (van Oosten, 1980a,b). It ding on established rootstocks, planted as lin-
also seems likely that Pseudomonas spp. bac- ers in the early spring. After the rootstock has
teria may be involved in the damage been cut back in the spring after budding, a
(Sholberg et al., 1993). shoot rises from the bud graft. Depending on
To prevent the ‘bleeding syndrome’, cut- the cultivar, it may or may not form laterals
ting back of rootstocks is best postponed (‘feathers’) (Plate 6.10). This formation of side-
until after the last spring frosts. However, shoots can be induced or improved using var-
this leads to smaller trees, because the ious techniques (see section on branching). In
remaining season available for their growth autumn, the trees are lifted and sold, either
is shorter. A compromise solution is to cut immediately or following an intermediate
back neither too early nor too late. In The storage period, outside or indoors. At the time
Netherlands, for those years that experience of sale, the tree head is 1 year old and the root
early bud development, the beginning of system 1 year older. Well-feathered 1-year-old
March is the recommended time and for trees have replaced the 2-year-old trees that
years with later bud break the second half of used to be the only branched trees available
March is preferred (van Oosten, 1980a). from nurseries. The latter were retained in the
Elsewhere pruning times may be different. nursery for a further year when no or few
Other measures considered beneficial, but side-shoots were formed in the first year. By
not proved, are to make a large wound or to cutting the vertical stem at heights varying
leave a rootstock stump (‘snag’) at the time from 85 to 110 cm, depending on the cultivar,
of cutting back. In the former case, a slanting branching was induced.
knife cut is made starting slightly higher Snip trees arise from rootstocks planted in
than bud height but at the opposite side of spring bearing a graft or a dormant (‘sleepy’)
the rootstock, ending about 5 cm above the bud. The scion buds grow out at the same
bud. In the second case, cutting back is done time as buds on conventional maiden trees,
at 10–15 cm above the bud. In both cases, a but in the first year development is moderate
larger surface area some distance from the because the roots have not established as well
as on the rootstocks used for the maiden trees, tree vigour. In environmental (soil and site)
which have been in the ground a year longer. conditions unsuited to the use of dwarfing
In the following winter, the single stem rootstocks, it is necessary to use tolerant and
(‘whip’) of the snip tree is cut at 50–80 cm often invigorating rootstocks to ensure tree
above the ground and only the highest bud is survival and longevity. This is relevant in
allowed to grow out. By then, the root system many parts of the southern hemisphere
is established and the terminal shoot grows where woolly aphid is a major pest of tree
out so vigorously that, depending on the cul- root systems. Dwarfing in these situations is
tivar, many laterals are usually formed (Plate achieved by using a dwarfing interstem,
6.11). Again, branching treatments may help such as M.9, together with a tolerant under-
to improve side-shoot formation. Currently, a stock, such as MM.106 or MM.111. These
cutting back height of 80 cm is preferred for interstock trees are raised using similar tech-
snip trees, especially with vigorous cultivars. niques to those described above.
This height gives a fairly long trunk, allowing The cropping level of snip and interstem
the fruiting laterals that develop from the lat- trees (Plate 6.12) is usually better than that of
erals on the tree following orchard planting to 1-year-old trees and for all tree types it holds
bend down under the fruit weight without that the higher the number of laterals, the
the need for premature shortening. In this higher the yield in the first years following
way, early cropping of difficult cultivars planting (Wertheim et al., 1995). For this rea-
becomes more feasible. The tree head of snip son, well-branched interstem or snip trees
trees is 2 years old. The name snip is derived are currently preferred by many apple grow-
from the Dutch word ‘knipboom’; ‘knip’ ers, even though they may be slightly more
means cut or snip, referring to the cut that is expensive to purchase. In areas where large
made to the 1-year-old stem. well-branched material would suffer from
Interstem trees are made using several spring droughts and where no irrigation is
methods. Traditionally, interstems are bud- possible, it might be better to plant less-
ded on to established rootstocks in late sum- developed material, which will establish
mer. Next year at approximately the same more easily in such conditions.
time, the scion cultivar is budded at about 50
cm above the ground on to the interstem.
Currently, interstem cultivars are also bud- 6.5.5 Tree support
ded on to bedded rootstocks in late summer
or bench-grafted on to bare-rooted rootstocks On dwarfing rootstocks tree support in the
in late winter. In both cases, the combina- nursery leads to the production of better
tions are spring-planted in the nursery for a trees. Bamboo canes or similar supports are
2-year production cycle. In the late summer placed into the ground next to each tree in
of the first season in the nursery, the scion the row at or shortly after bud break. The
cultivar is budded at about 50 cm above main shoot is regularly attached to the sup-
ground level on to the interstem. With the ports with tape, usually using a special tying
lower rootstock union at 15 cm, the interstem apparatus. The support prevents leaning or
is usually 35 cm long, but its length may breakage at the union and contributes to a
vary according to local requirements. high growth rate of the shoot leader, an
Interstems are used to provide the tree- essential condition for feathering (Tromp
trunks with resistance to diseases (the Dutch and Boertjes, 1996).
apple cultivar ‘Dubbele Zoete Aagt’ against a
trunk rot caused by Phytophthora cactorum
(Lebert & Cohn) Schroter) or to freeze dam- 6.5.6 Trunk cleaning
age (‘Hibernal’ or ‘Summerred’). Moreover,
interstem trees branch as freely as snip trees Shoots may arise from the trunk in positions
and they similarly raise the height of the where they are not wanted. In the final year
fruiting laterals. In many parts of the world of the growth of 1-year-old trees, shoots orig-
interstems or interstocks are used to control inating lower than 40–50 cm above the soil
are unwanted. If retained and allowed to buds), stored, planted out in the nursery in
develop, fruits borne on such branches hang the following spring and raised as snip trees
too close to the ground and are thus difficult during the next two seasons. Bench grafts
to pick and prone to rot because soil fungi that are made indoors during the late
may splash up with soil particles during winter are planted out in spring and the
rain. Laterals that are too low are rubbed trees also develop during the next two
away when still only a few centimetres long. growing seasons, as for snip trees. The
By doing this early, wounds are small and longer period needed for snip trees is due to
heal easily. This measure may slightly favour them making only moderate scion develop-
the formation of higher laterals and has no ment (‘whips’) in the first year. This is
negative effects on trunk size or tree height because the rootstocks are not sufficiently
(de Groene, 1986). With snip and interstem established in the nursery in the first spring.
tree raising, shoots below the preferred ter- Budding makes more economical use of
minal shoot are also rubbed away, but in scion propagation material than grafting
stages, with time intervals between each and is more popular currently than grafting.
removal in spring and early summer, to pre- Where rootstocks have failed to reach suffi-
vent bleeding. cient size for budding in the previous sum-
mer, trees are occasionally grafted in early
spring outside in the nursery. In the past,
6.5.7 Other types of plant material when buds that were inserted in rootstocks
during the summer failed to ‘take’ (failed to
Apple-tree raising in containers is not con- heal and form a viable union with the root-
sidered an economically viable option, stock) they were grafted in the subsequent
because it requires a great deal of attention spring. This is not recommended, however,
to detail and is expensive and establishment as poor bud take is usually indicative of
is more difficult (Wertheim and de Groene, graft incompatibility, virus infection or
1988; Wertheim and Wijsmuller, 1988; some other health problem with the stock or
Wertheim, 1989). Similarly, planting of ‘half- the scion.
finished’ products directly in the orchard Budding and grafting are also used in
cannot be recommended. Bench grafts or top-working established trees in the orchard.
rootstocks with dormant scion buds give dis- Top grafting (also termed top working or
appointing yields compared with finished frame working) has recently become quite
trees and result in more tree losses and tree- popular in the UK as a method of rapidly
to-tree variability (Bootsma and Baart, 1990). converting an existing orchard over to one or
more new scion cultivars. Significant yields
are produced several years in advance of
6.6 Budding and Grafting planting new trees using this technique and
the costs of converting the orchard are much
In commercial nurseries, apple trees are less than when purchasing new trees. The
most often produced by either late-summer technique is only to be recommended on
budding of scion cultivars on to rootstocks trees that were known to be virus-free at the
in the field or by bench grafting them on to time of planting the original orchard. Unlike
bare-rooted rootstocks indoors in late win- stone fruits, where viruses are transmitted
ter. Two systems of budding may be used to easily in pollen, apple viruses seem to be
raise trees. In the first, budding is done on transmitted only by budding and grafting,
to established rootstocks planted in the pre- and healthy orchards generally remain clean
ceding dormant season and in the following throughout their lives. Top- and frame-work
year the bud develops into a 1-year-old tree grafting of apple trees is carried out during
(‘maiden tree’). In the second system, bud- the dormant season using healthy, virus-free
ding is done on to closely planted bedded scion wood. Details of the various tech-
rootstocks, which are then dug up in niques used are given in Garner (1979) and
autumn (with what are called ‘sleepy’ Hartmann et al. (1990).
6.6.1 Bud wood rooting later in the orchard, but 15 cm is a

better standard. For special purposes, such
In budding, a piece of stem and bark of a as increasing dwarfing or alleviating prob-
scion cultivar carrying one bud is inserted lems of disease, budding height is higher.
into the stem/bark of a rootstock. Bud wood For growth control of vigorous scion culti-
is taken from the middle part of current-year vars, even on dwarfing rootstocks, such as
shoots obtained from mother trees of a certi- M.9, heights of 25–35 cm are often preferred.
fied healthy source. Shoots used should pos- Also, with ‘Cox’s Orange Pippin’, a cultivar
sess healthy leaves and well-developed very susceptible to Phytophthora trunk rot,
buds. This rules out the use of the succulent budding height is at least 30 cm. In this way,
shoot tip as well as the woody basal part. the soil-borne fungus cannot reach the trunk
Shoots with flower buds are discarded. After of the scion cultivar by the splashing of soil
collection, all leaf blades and stipules are particles containing the pathogen during
quickly removed to prevent dehydration. heavy rains.
Leaf petioles are usually left intact at this It is usually irrelevant which side (orien-
deleafing to facilitate bud insertion and as a tation) the bud is inserted into the rootstock
later indicator of success or failure of bud stem, but in windy conditions budding on
take. With successful bud take the petiole the windward side reduces chances of the
drops off, whereas it withers and stays on buds being pushed out by strong winds.
where a bud does not unite with the root- However, when shoots are tied to support-
stock. However, petioles can also be ing bamboo canes, this is not necessary.
removed with the blades, because the above When budding is done on over-row carts, to
advantage is considered of minor impor- make the work less tiring for personnel,
tance by some nurserymen. buds are inserted in the same direction as
After leaf removal, the shoots of bud the row because the rootstocks are bent over
wood should be used as soon as possible. In that way.
cool and moist conditions, they can be stored
for up to 1 week, but it is better to always
use freshly harvested material where possi- 6.6.3 Budding and grafting methods
ble. As long as well-developed buds are
used, the bud origin on the shoot is of no Two techniques of budding, T- and chip bud-
consequence for bud take or tree quality ding, are used for propagation of apple trees.
(Smith et al., 1962). To smooth peaks in In both techniques it is essential to achieve
demand for labour needed for budding, bud effective joining of the cambial layers of the
wood can be stored for up to 4 weeks at 2°C. scion bud and the rootstock and to prevent
Longer storage gives lower bud take desiccation.
(Versteegen, 1989). In the field, bud sticks In T-budding, a T-shaped incision is made
should be kept fresh by keeping them moist in the rootstock rind at the time in summer
and cool. when the bark easily slips from the underly-
ing xylem to facilitate bud insertion. In the
northern hemisphere this happens from late
6.6.2 Rootstocks, budding height and site July till early September. An upper trans-
verse cut is made first, about 1 cm long. A
To be suitable for budding, rootstocks second, vertical cut of about 2.5 cm is made
should be neither too thick nor too thin. In upwards, such that it meets the first one at
the case of M.9, suitable basal-diameter its middle point. The bud graft is a thin, nar-
classes at planting are 5–7 mm and 7–9 mm row, oval-shaped piece of bark including a
(van Oosten and de Groene, 1980). central bud and with some wood tissue on
Preferably, the two classes are planted in the inside. Holding the bud stick by the
separate blocks as they may render trees of upper end a cut is made beginning about 1
different quality. Budding is done at least 10 cm below a bud. Thereafter, the knife is
cm above ground level to prevent scion moved shallowly upwards beneath the bud
to about 2.5 cm above it. The shield is then canker may develop. However, in hot areas
torn from the stick leaving a bark strip that T-budding might be the better procedure,
facilitates insertion. After insertion, this strip because the protective bark slips decrease
is cut away and the two rootstock bark flaps the risks of desiccation (Hartmann et al.,
are bent back over the bud graft (Anon., 1990). Desiccation is the major problem with
1963; Hartmann et al., 1990). chip budding and it is only since the advent
In chip budding, an oval shield with the of polythene or rubber tying materials that
central scion bud replaces an exactly simi- the popularity of the technique has
larly shaped piece of bark removed from the increased; chip budding was rarely success-
rootstock and consequently the cambia of the ful when only raffia bud ties were available.
two partners usually match very well (Plate Commercially, both methods of budding are
6.13). Both chips are cut out in a similar man- currently used successfully. The fate of bud
ner. On the bud stick, the first cut is made 0.5 take can be non-destructively determined by
cm below the bud down into the wood at an magnetic resonance imaging (Warmund et
angle of 30–45° to a depth of approximately al., 1993) – useful in cases of new scion–root-
one-quarter of the rootstock diameter. A sec- stock combinations where graft incompati-
ond cut starts about 2.5 cm above the bud bility may be suspected.
and goes inwards and downwards behind it
until it meets the first cut. The sequence of
making the cuts may be reversed. In the 6.6.4 Tying and after care
rootstock a similar chip is removed in a simi-
lar manner. The cambia of bud stick and Tying must be done very soon after bud-
rootstock should be opposite each other, at ding, either with rubber bands, which dete-
least at one side of the union. The good jux- riorate (break down) after a few weeks so
taposition of the cambia of scion and root- that the thickening rootstocks are not gir-
stock in chip budding leads to a rapid fusion dled, or alternatively with 1 cm wide plastic
and good bud take. For chip budding it is strips of polyvinyl chloride (PVC) film (Plate
not strictly necessary for the rootstock bark 6.14). This latter material is elastic, moisture-
to slip easily at the time of budding; this proof and transparent, allowing inspection,
gives slightly greater flexibility and a longer but it does need to be cut after the buds
time period during which budding can be have healed. This is done with a superficial
undertaken. knife cut made to the plastic on the side
In lifting the bark from the rootstock in T- opposite to the bud. Rapid and complete
budding, the cambium zone remains cover of the wound and bud may help to
attached to the inside of the bark. Therefore, prevent damage by the red-borer fly
the rootstock cambium cannot quickly match (Thomasiniana oculiperda Rubs). In areas
with the scion-bud cambium (Mosse and where this pest is not a problem, other ties
Labern, 1960). This has consequences for may need to be used.
their coalescence and may affect bud take It is very important that the binding
and tree quality adversely. For these reasons, material does not girdle the rootstock, as this
chip budding seems the better method. will delay growth and lower tree quality. For
Compared with T-budding, chip budding the same reason, budding should not be
often resulted in better bud take, less freeze done too early in the season when much
damage of buds and more developed and rootstock stem thickening still has to occur.
uniform trees (Howard, 1974; Howard et al., Moreover, early budding can stimulate the
1974; Skene et al., 1983). However, in a few bud grafts to grow out shortly afterwards.
studies no benefits of chip budding have This is unwanted as the little shoots can eas-
been shown (Meiß, 1985). A minor drawback ily be damaged by winter freezes. In the
of T-budding is that spores of the fungus northern hemisphere, most thickening
Nectria galligena (Bres.), which may be pre- occurs in July and August. Hence, budding
sent on the outside of the bud graft, can be in the second half of August is better than in
introduced into the rootstock and fruit-tree July (Smith et al., 1962).
After budding, the rootstock is not shoot growth (SSG). Controlled-environment

pruned until next spring to keep the bud experiments have shown that an 18-week
graft dormant. In spring, the rootstock part period with air temperatures of 25°C, start-
situated above the bud is cut off either coming at bud break, greatly enhances SSG, com-
pletely or in stages. pared with a similar period at 15°C. When a
temperature of 25°C was given for 6 weeks
following a similar period with low tempera-
6.6.5 Bench grafting tures, SSG was also greatly favoured,
whereas 15°C in the second 6-week period
Bench grafts are made indoors during the had no effect (Tromp and Boertjes, 1996). The
winter season. The main method used is the higher the soil temperature in the range
whip-and-tongue graft (Garner, 1979; 7–28°C during the growing season, the more
Hartmann et al., 1990). In order to economize laterals are formed (Tromp, 1992a). SSG was
on costs for propagation material, the grafts strong when soil temperature was 12°C dur-
are short, bearing only two buds. With whip- ing the first 6 weeks after bud break fol-
and-tongue grafting, the diameter of the lowed by a similar period of 22°C (Tromp,
stem of the scion and the rootstock should be 1996). High humidity also favours lateral for-
about equal. When the rootstock is thicker, mation (Tromp, 1992b). A high humidity
side-grafting is used, although it is known (90%) given in the first 6 weeks after bud
that the union of the two partners is less break promoted SSG at 22°C soil tempera-
strong than with a whip-and-tongue graft. ture, but not at 12°C (Tromp, 1996). The
After grafting, the two partners are firmly interaction of external factors means that
tied together with a plastic strip and the api- feather formation in the nursery will vary
cal end of the graft is sealed with grafting depending upon seasonal conditions. It also
wax. Bench grafts can be planted out directly follows that some climatic conditions or sites
but, when soil conditions do not permit this, are more favourable for the raising of feath-
storage at about 2°C under high humidity, to ered trees than others.
prevent dessication, is necessary. It has long been assumed that the apex of
the parent shoot inhibits lateral growth by
auxins produced in the tip, which are then
6.7 Branching transported basipetally. However, the back-
ground to lateral-shoot inhibition is more
The vigorously growing vertical shoot of the complex than this (Cline, 1994). It is possi-
apple scion (the central leader) that arises ble that competition for water and nutrients
from the bud, may or may not form sylleptic is involved, for when these are limiting only
shoots, or ‘feathers’. For a high-quality tree, the main shoot grows. Whatever the cause,
the presence of a good number of feathers is SSG in apple mainly occurs when the
desirable, because they form flower buds in growth rate of the main shoot is highest
the year following orchard planting and (Tromp, 1996).
enable the tree to bear fruit in the second year.
The more feathers, the higher the yield in the
first years (van Oosten, 1978; Wertheim, 1981), 6.7.1 Manipulation of branching by hand
and all nursery efforts should be directed
towards raising feathered trees. Removal of the growing tip induces lateral-
Cultivars vary greatly in their tendency to shoot formation, but tipping is not a good
form sylleptic shoots. Some hardly feather at technique, because it induces laterals that
all, while others do so very freely. There is grow out with very narrow branch angles
also a year-to-year effect on feathering, (Cody et al., 1985; Jarassamrit, 1989). In the
pointing to a role of environmental condi- orchard, narrow-angled branches can break
tions in the induction and development of off easily under a heavy fruit load and nar-
feathers. Indeed, air and soil temperature row crotch angles may also become infected
and humidity have great effects on sylleptic by N. galligena (Bres.), leading to fruit-tree
canker. Narrow angles also arise when the After tipping and removal of the apical
growing tip is killed by use of too aggressive meristem, shoots arise under the wound, but
chemical branching agents. In contrast, when with leaf removal as described above they
a shoot tip is gently reduced in growth activ- arise both below and above the height of
ity, well-positioned laterals with broad treatment (Wertheim, 1978a,b). Taking away
branch angles arise. This gradual slowing leaf blades can easily and quickly be carried
down of the speed of growth of the leader out by casual labour. In the case of some
can be achieved manually by removing the difficult-to-feather cultivars, some nurserymen
leaf blades from the actively growing tip, deleaf up to six times in a growing season.
leaving the actual growing point intact. A sin-
gle removal of leaf blades exerts an effect, but
repetitive treatments at 7–14-day intervals are 6.7.2 Manipulation of branching by
more effective (Table 6.4). It is important that chemicals
all young leaves that are still light green in
colour are removed completely. Removal of A chemical branching agent applied once
only parts of the young leaves is not effective can have a beneficial effect on feathering
(Wertheim, 1978a), nor is taking away mature (Table 6.4), but mixtures applied once or
leaves under the shoot apex (Table 6.4). The repeated sprays with the same compound or
most effective treatment of the trial summa- with two different chemicals are usually
rized in Table 6.4 was a combination of three more effective. Effective chemicals are: ‘M&B
successive removals of young apical leaves 25,105’ (propyl 3-t-butylphenoxy acetate),
plus a ‘Promalin’ spray. This produced 8.4 BA, GA4+7 (gibberellins A4 + A7), and a mix-
feathers per tree compared with 2.6 for the ture of 50% each of BA and GA4+7, such as
untreated control. ‘Promalin’. All chemicals act better when
Table 6.4. Results of a branching trial with ‘Red Boskoop’ on M.9 rootstock. Sixteen treatments were
compared, namely, one spray of ‘Promalin’ when the leader was 65 cm in height at 1000 p.p.m. each of
BA and GA4+7 (P); stripping the apex one, two or three times at 10-day intervals of all young leaves
(commencing when leader height 65 cm) (A); and stripping the sub-apical zone only once (45–65 cm
above the ground) when the leader height was 70 cm (SA) and all combinations (data not shown).
(Adapted from Wertheim et al., 1989.)
Tree height Stem diameter Feathers per Average feather

Treatmenta above union (cm) (mm)b tree 10 cm (no.) length (cm)
−P 139.5 14.2 4.0 22.1

+P 136.4 14.6 7.3 24.2
F test NS NS *** NS
LSD0.05 – – 0.7 –
−SA 137.4 14.4 5.7 22.4
+SA 138.6 14.5 5.6 23.9
F test NS NS NS NS
LSD0.05 – – – –
A 0× 139.9 14.1 4.7 15.9
A 1× 136.4 14.1 5.5 21.6
A 2× 137.9 14.6 5.9 27.5
A 3× 137.6 14.8 6.7 27.6
F test NS NS *** *
LSD0.05 – – 0.9 8.8
aSee table title.
b20 cm above union.
F-test interactions P × SA, P × A and A × SA not significant; thus 16 treatments combined.
NS, not significant; *, significant (P 0.05); ***, very strongly significant (P 0.001).
applied with a wetting agent (Wertheim, raised – for example, from 4.0 in control trees
1978a,b; Gerhard, 1984; Cody et al., 1985; to 15.9 per tree after eight weekly sprays
Jarassamrit, 1989; Wertheim et al., 1989; Volz with 400 p.p.m. BA (Wertheim and de
et al., 1994; Wertheim and Estabrooks, 1994). Groene, 1993). Results differ per cultivar; in
As with manual treatments, branching one nursery where four or six sequential
agents only have their beneficial effects if weekly sprays with 150, 300 or 600 p.p.m.
applied to a vigorously growing shoot. It is BA were evaluated, ‘Elstar’ reacted best to
sufficient to treat only the top 25 cm of the four sprays of 300 p.p.m. BA, ‘Cox’s Orange
shoot, although ‘Promalin’ also acts when Pippin’ to four times 300–600 p.p.m.,
applied to mature leaves and to lateral buds ‘Delcorf ’, ‘Golden Delicious’ and ‘Jonagold’
(Gerhard, 1984). Applications can start at a to six sprays with 300 p.p.m., and ‘Red
leader height of 45 cm, but this may induce Boskoop’ to six sprays with 600 p.p.m. With
laterals to form too low on the leader, for all the cultivars, spectacular increases in
they arise a few decimetres below and above numbers of laterals were obtained and sub-
the height of treatment (Wertheim, 1978a). sequent orchard performance was not nega-
After eight weekly sprays with 200–400 tively affected (Wertheim and de Groene,
p.p.m. BA treatments on ‘Red Boskoop’, 1995). Eight sprays of 200–400 p.p.m. BA
starting when the leader was 35 cm above may diminish the flower-bud formation that
the budding height (50 cm above the sometimes occurs on 1-year-old trees
ground), the zone of lateral emergence was (Wertheim and Estabrooks, 1994). This may
enlarged from 31–70 cm to 31–120 cm above be regarded as an advantage, as the trees can
the union (Wertheim and Estabrooks, 1994) establish more easily without fruits being
(see Plate 6.10). With single sprays, best formed in the planting year. An example of
results are obtained when shoot tips are the possibilities of using a series of BA
approximately 65–70 cm above ground level sprays is presented in Table 6.5. A combina-
at the time of treatment. Sequential sprays tion of repeated manual deleafing and one
may begin at a lower leader height – for ‘Promalin’ spray proved slightly more effec-
example, 50 cm. If the growth level is insuffi- tive than ‘Promalin’ alone (de Groene, 1990).
cient, only thick lateral buds may arise, but For success in branching, not only the
no lateral shoots. Although the treatments do vigour of growth is important; the environ-
not affect the trunk size, they may reduce ment must also be favourable. Weather fac-
final tree height, but this effect is normally tors cannot be influenced, but other factors
not of commercial importance. can. Only soils of excellent quality should
Successful chemical treatments are be chosen, rootstocks at planting time
achieved using 500–1000 p.p.m. ‘M&B 25,105’ should be of adequate size and within-row
followed 2 weeks later by 4% ‘Promalin’ (= planting distances ample to allow for good
720 p.p.m. of both BA and GA4+7) (Gerhard, lateral growth (Wilson and Jarassamrit,
1984). Alternatively, a mixture of 1000 p.p.m. 1994). At a planting distance of 100 cm × 10
‘M&B 25,105’ + 250–500 p.p.m. ‘Promalin’, or cm, feathering of several apple cultivars
a spray of 1000 p.p.m. ‘M&B 25,105’ fol- was rather poor, although improved by BA
lowed 1 week later by 500 p.p.m. ‘Promalin’ and GA4+7 combinations and sequences
(Cody et al., 1985) can also be effective. (Volz et al., 1994).
Mixtures of 1000 p.p.m. BA and 1000 p.p.m. There is a reluctance to use branching
GA4+7 or sequences of these two hormones agents among nurserymen, because of incon-
with 2-week intervals are also successful sistent results and/or lack of demand for
(Volz et al., 1994). Also very successful were feathered trees (Miller, 1988). Given the
one or two sequences of BA and GA4+7, each effects of environmental factors, inconsis-
at 1000 p.p.m., with 2-week intervals, as tency is to be expected, although this can be
were two successive applications of reduced by repeated treatments (Table 6.5).
‘Promalin’ with a 2-week interval between, Reluctance to induce feathering may also be
or 1000 p.p.m. of both hormones. The num- based upon the more difficult handling of
ber of feathers is sometimes spectacularly feathered trees during harvest, storage and
Table 6.5. Results of a branch induction trial with ‘Elstar’ on M.9 rootstock. The first ‘Promalin’ spray was
applied when the central leader was 70 cm high (55 cm above the union) at 1000 p.p.m. of BA and
GA4+7. The second spray was applied 2 weeks later. The BA spray was applied one, two, four or eight
times at weekly intervals at the concentrations shown.
Tree height Feathers per Spurs per Average feather Total per
from union tree 10 cm tree 10 cm length treeb
Treatmenta (cm) (no.) (no.) (cm) (cm)
Control 127.6 4.3 1.6 32.8 284.8

1× ‘Promalin’ 131.5 6.9 1.6 37.8 389.8
2× ‘Promalin’ 127.5 6.5 2.4 40.5 391.8
1× 200 p.p.m. BA 131.1 6.5 1.6 37.5 368.1

2× 200 p.p.m. BA 125.0 6.1 1.6 38.9 365.5
4× 200 p.p.m. BA 125.5 6.5 1.3 46.8 435.3
8× 200 p.p.m. BA 125.4 7.4 1.4 43.6 444.5
1× 400 p.p.m. BA 124.9 5.1 2.9 44.2 360.4

2× 400 p.p.m. BA 129.5 6.0 0.9 43.4 396.0
4× 400 p.p.m. BA 125.3 7.1 1.8 42.0 432.6
8× 400 p.p.m. BA 121.0 9.0 2.6 38.3 471.9
1× 800 p.p.m. BA 129.3 6.4 2.3 41.9 410.4

2× 800 p.p.m. BA 127,5 7.9 0.5 44.7 478.0
4× 800 p.p.m. BA 119.6 7.6 3.0 36.5 403.5
8× 800 p.p.m. BA 124.3 13.0 3.3 37.7 617.1
F treatment ** *** ** ** ***

LSD0.05 5.4 1.7 1.3 6.4 71.9
aApplied to 15 cm apex.
bFeathers + spurs + tree height.
**, Strongly significant (P 0.01); *** very strongly significant (P 0.001).
transport. Although an understandable reac- are damaged after lifting. In areas where
tion by the nurseryman, fruit growers should cold nights in early autumn prevail, natural
insist on the supply of well-feathered trees. leaf abscission occurs satisfactorily with
With snip and interstem trees the chemi- most cultivars. However, in maritime cli-
cal approach to feathering is more difficult, mates with mild autumns coupled with rela-
since, at the application height used with 1- tively warm nights, some cultivars do not
year-old trees, the young shoots are still too shed their leaves easily. In these cases, labori-
short and can be easily damaged (Wertheim, ous hand defoliation is needed before trees
1986b). In this instance, applications, if nec- are lifted. Many efforts have been made to
essary, should be made later when the shoots find safe chemical defoliants. Ideally, they
are still actively growing but also have a suf- should not damage the tree and not
ficient number of mature leaves. adversely affect growth in the next season
(Miller, 1988). Ethephon, for example, can
induce leaf abscission in apple but with the
6.8. Defoliation And Digging Up concentrations needed, growth in the plant-
(Lifting) Trees ing year may be reduced, and this is highly
undesirable. Copper chelate has in recent
It is customary to dig up apple trees from the years emerged as a reliable defoliant for
nursery in autumn, after all shoot growth apple. Depending on the cultivar, one or two
has stopped and most leaves have abscised. sprays with 1–2% of a 9% copper-chelate
Trees with leaves still present desiccate and product plus a wetting agent, applied at the
time that some signs of autumn leaf discol- root systems are laid in an opposite direc-
oration and/or some natural leaf drop has tion. A few people can lift 1 ha of trees day−1,
begun, induce good defoliation without any even from heavy, moist, clay soil. Tree han-
subsequent damage. In north-western dling during lifting should proceed carefully
Europe, this means an application some- to prevent breakage of laterals. After lifting,
where between mid-September and the end trees can either be shipped directly to cus-
of October (Wertheim, 1984; Faby, 1989). tomers or stored until delivery in the spring.
However, its use is only possible in countries Trees can be clamped (‘heeled in’) outside in
where it is registered (approved) for use. In bundles situated in a cool, sheltered place
the USA, a combination of ethephon (150 mg with their roots well buried into the soil and
l−1), the surfactant ‘Depeg’ (0.5%) and protected against rodents by chicken-wire. A
‘Alanap’, the sodium salt of N-1-naphthylph- safer option is to store the trees indoors in
thalamic acid (200 mg l−1), applied in rooms with high humidity at a few degrees
October, showed promise in defoliation stud- above 0°C. Under no circumstances should
ies with apple, but this mixture requires fur- trees be stored in rooms containing ethylene
ther testing before it can be widely in the air – for example, emanating from fruit
recommended (Larsen and Higgins, 1999). or residual ethylene impregnating store
Tree lifting from the nursery, which walls after fruit storage – as this leads to
involves undercutting the trees, has been reduced bud break on the trees in the spring.
largely mechanized, for which tractor-drawn
or self-driving machines are used. The trees
are undercut with a u-formed knife, lifted Note
from the soil by metal fingers, clasped
between two upward-moving rubber con- Neither the authors nor their research institutes
veyor belts, which shake to remove the soil can accept any liability for loss, damage or
from the roots, and transported to a pallet, injury resulting from the application of any con-
where people lay them carefully in a hori- cept or procedure in or derived from any part of
zontal position. With each layer of trees the this chapter.
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Wertheim, S.J. and Kunneman, B.P.A.M. (1993) Onderzoek naar M.26 in de fruitteeltfase. Onderstammen
uit weefselkweek zijn bruikbaar. Fruitteelt 83(45), 20–21.
Wertheim, S.J. and Wijsmuller, J. (1988) Plukken in het plantjaar. Fruitteelt 78(21), 16–17.
Wertheim, S.J., de Groene, J.M. and van de Wassenberg, J. (1989) Promalin plus pluizen geeft meer veren.
Fruitteelt 79(21), 18–19.
Wertheim, S.J., Bootsma, J.H. and Jansen, W.A.G.M. (1995) Knip- en tussenstambomen voldoen beter dan
eenjarige bomen. Fruitteelt 85(28), 14–15.
Wilson, S.J. and Jarassamrit, N. (1994) Nursery factors influencing lateral shoot development in a spur
type apple cultivar. Scientia Horticulturae 56, 207–215.
Zhu, L.H. and Welander, M. (1999) Growth characteristics of apple cultivar Gravenstein plants grafted on
to the transformed rootstock M.26 with rolA and rolB genes under non-limiting nutrient conditions.
Plant Science 147, 75–80.
Zimmerman, R.H. (1997) Orchard variation in micropropagated trees of ‘Redspur Delicious’ apple.
7 Flowering, Pollination and Fruit Set and

Development
Frank Dennis, Jr
Department of Horticulture, Michigan State University, East Lansing, Michigan, USA
7.1 Juvenility 153

7.2 Flower Induction, Initiation and Development 154
7.3 Flowering Habit, Flower Structure 157
7.4 Pollination and Fertilization 157
7.4.1 Parthenocarpy 157
7.4.2 Pollination 157
7.4.3 Fertilization 160
7.5 Fruit Development 160
7.6 Seed and Fruit Growth 161
7.7 Fruit Maturation 162
7.8 Summary 164
Flowers are the ‘raw materials’ for fruit pro- years, depending upon species. This creates a
duction. Therefore heavy flowering is essen- problem for apple breeders, who often must
tial for economic success. But, unlike the wait for 7–9 years before being able to see the
situation with floriculture crops, flowers fruits of their labours. Treatments that stimu-
alone are not sufficient. They must set fruit late flowering in mature tissues, such as ring-
and the fruit must mature and be of suffi- ing of the bark (see below), have little or no
cient size and quality to be marketed at a effect on seedlings until they reach ‘ripeness
profit. This chapter describes the flowering to flower’. Grafting scions from seedling trees
process in apple, how flowers develop into into bearing trees can hasten flowering, but
fruits, and how fruits grow and mature. the time gained is only a few years. Another
method used by breeders is growing the trees
as single stems. Flowering can be induced in
7.1 Juvenility the second year in some cases, but only buds
near the top of the tree flower; there appears
Seedlings, whether of herbaceous or woody to be an effect of ‘distance from the roots’,
plants, must pass through a juvenile period suggesting that inhibitory compounds (gib-
before they are capable of flowering. In berellins (GAs) ?) produced in the roots move
herbaceous plants this may require a few up the stem and prevent flowering. If the dis-
weeks; in most trees it requires from 1 to 20 tance between roots and buds is sufficient,

154 F. Dennis, Jr
the concentration of these compounds is too 7.2 Flower Induction, Initiation and
low to be inhibitory. This hypothesis is ques- Development
tionable, however, because buds taken from a
juvenile tree will not flower when grafted Flower induction refers to the change from
into the top portion of a bearing tree. vegetative to reproductive phase and can be
Commercial cultivars are ‘mature’ (no likened to a switch. However, no visible
longer juvenile); the original seedlings were macroscopic or microscopic changes occur in
juvenile, but bud wood taken from the the bud. Most flower induction occurs in
mature, upper parts of these seedlings has early summer, but it can extend into early
been used for propagation through many autumn under some conditions. Initiation
‘generations’ of trees. Young trees on seedling begins when the meristem flattens – visible
rootstocks generally require several years microscopically – and continues as primor-
before they form flower-buds. The time re- dial sepals, petals, stamens and pistils form
quired varies with cultivar, ‘Golden Delicious’ centripetally on the apex and grow into fully
being very precocious and ‘Northern Spy’ formed appendages (Fig. 7.1). Most of the
very late in this respect. When young trees are flower parts are present by early autumn,
very vigorous, flowering tends to be delayed. but continue to develop in temperate cli-
Propagation on dwarfing rootstocks reduces mates until low temperatures prevent fur-
the time to flower considerably, with most ther growth. In such climates, meiosis begins
dwarf trees flowering within 2–3 years. in the anthers in late winter.
a b
c d
Fig. 7.1. Stages in the initiation of an apple flower at Long Ashton, Bristol, UK, 1974 (Abbott, 1977).
Scanning electron-microscope views. (a) 1 August – vegetative apex with five leaf primordia; (b) 15 August –
apex becoming domed, ridges (floral initials) arising in leaf axil/bract primordia; (c) 21 August – longitudinal
section showing ‘king’ flower at apex; (d) 19 September – five sepals forming on the uppermost lateral
flower initial (subtending leaves and ‘king’ flower removed).
Flowering, Pollination and Fruit Set and Development 155
In most fruit-growing areas buds become if vigour is excessive, the optimum number
dormant in late summer/early autumn and is exceeded; if vigour is too low, too few
winter chilling is necessary to permit nodes are formed and, in either case, shoots
renewed growth the following year (see fail to form flower buds.
Chapter 10). Weinberger (1950), working in Flower induction can be inhibited by
California, used the cumulative number of heavy cropping, some cultivars (e.g. ‘Yellow
hours at or below 45°F (= 7.2°C) as an indica- Newtown’, ‘Paulared’, ‘Fuji’) being notori-
tion of the amount of chilling received by ous for their ‘biennial bearing’ habit,
peach trees. Although this model has been although all cultivars exhibit some degree of
used at higher latitudes, temperatures below response to heavy cropping. This can be con-
freezing have very little or no effect in break- trolled in most cultivars by early fruit
ing dormancy. A model developed by removal (‘thinning’), either by hand or with
researchers at Utah State University (the chemicals (see also Chapter 16). In general,
‘Utah model’) can be used to predict fruit must be removed within the first 3 or 4
response in areas with colder winters weeks following bloom for thinning to be
(Richardson et al., 1974). Temperatures effective, response declining as thinning is
between 0 and 15°C are effective, with maxi- delayed (Fig. 7.2). The physiological basis for
mum response at 6–7°C, where 1 h of chill- this effect of fruits on flowering has been the
ing equals 1 ‘chill unit’ (CU). Temperatures subject of much research, but remains to be
16°C have detrimental effects, reducing determined. Apple seeds are rich sources of
the response to previous chilling. This model GAs. These compounds can inhibit flower-
works well in the temperate zone, but is less ing when applied to limbs or whole trees,
useful in the subtropics. In these warmer and most of the theories proposed include a
areas the ‘dynamic’ model (Erez et al., 1988) role for them in inhibiting flowering. In the
is a better predictor of response. This model facultatively parthenocarpic apple cultivar
assumes that chilling is accumulated in units ‘Spencer Seedless’, seeded fruits inhibit flow-
and, once such a unit is acquired, high tem- ering, whereas seedless fruits do not, sug-
perature cannot nullify its effect. gesting that seeds do play a crucial role in
Cultivars differ in the amount of chilling flowering (Chan and Cain, 1967; Neilsen and
necessary to break dormancy. Some (e.g. Dennis, 2000). However, few such cultivars
‘Anna’, ‘Dorsett Golden’) require as little as exist, and the evidence for the role of seeds
250–300 CU and can be grown in the sub- in related species, such as pear (Pyrus com-
tropics, whereas others (e.g. ‘McIntosh’) munis L.), is more controversial (see Dennis
require much more chilling (1000–1600 CU) and Neilsen, 1999; Weinbaum et al., 2001).
and can only be grown at higher latitudes of
the temperate zone. If chilling is insuffi-
cient, both vegetative and flower buds are 50
Spurs forming blossoms (%)
retarded in development and cropping is

reduced. Exceptions to the requirement for 40
chilling occur in some regions of the tropics,
as in Indonesia, where defoliation soon 30
after harvest induces bud break, resulting in
two crops per year (Edwards and 20
Notodimedjo, 1987).
Abbott (1977) reported that 20–24 nodes 10
must develop before apple flowers can be
initiated. Although studies with potted trees 0
had suggested that flower ‘quality’ (= ability 20 30 40 50 60 70 80
to set fruit) was optimum when initiation Days after full bloom
occurred at a specific time, Abbott observed Fig. 7.2. Effect of time of thinning ‘Yellow Newtown’
little difference in time of initiation under apple fruits, to leave one fruit per 70 leaves, on
field conditions. Others have suggested that, flowering the following year (Harley et al., 1942).
156 F. Dennis, Jr
Environmental factors also affect induc- of some cultivars (e.g. ‘Empire’) have wide
tion and initiation. Although photoperiod crotch angles, which permit better light pene-
plays little or no role in flowering of apple tration; others (e.g. ‘Delicious’) have narrow
under field conditions, solar radiation is crotch angles. Leaves, of course, are important
important. Flowering is heavier in well- for capturing sunlight, and defoliation
exposed sections of the tree and in trees in reduces flower induction/initiation. Leaf
areas with high solar radiation, such as injury from insects and disease can therefore
Washington State and California. reduce flowering. In controlled environments,
Experiments with artificial shading have high temperature can inhibit flowering in
indicated that flowering is reduced when- some cultivars (Tromp, 1976), but evidence
ever the light level is reduced below 30% of that this occurs under field conditions in the
full sun (Fig. 7.3). Pruning to open the tree to temperate zone appears to be lacking.
sunlight is therefore encouraged. Young trees Many cultural practices, including ring-
ing or scoring the trunk or scaffold limbs and
bending of limbs, favour flower induction.
80
Ringing/scoring is recommended for young
trees only when flowering is delayed consid-
erably, as in ‘Northern Spy’, but bending or
70 spreading limbs to increase exposure to light
is a common practice in young orchards (see
Fruit retained (%)
Chapter 14). Drought conditions tend to

60 favour flower-bud formation, although this is
not well documented and some researchers
50
have questioned the relationship. Heavy
applications of nitrogen stimulate growth
and tend to reduce flower-bud formation.
40 However, limited evidence exists that appro-
priate timing in applying summer nitrogen
can improve flower quality, leading to better
30 fruit set (Williams, 1965; Hill-Cottingham
25 50 75 100
and Williams, 1967).
Some plant growth regulators affect flow-
ering. Tri-iodobenzoic acid can promote
180
flowering in ‘Delicious’. Several other
compounds, including ethephon (2-
chloroethylphosphonic acid) and the growth
Mean fruit weight (g)
retardants butanedioic acid mono-(2,2-

dimethylhydrazide) (Alar®), 2-chloroethyl
trimethyl ammonium chloride (CCC), and
160 paclobutrazol ((1RS,3RS)-1–1-(4-chlorophenyl)-
4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-pentan-
3-ol), are effective on numerous cultivars,
although neither Alar® nor paclobutrazol can
be used commercially in the USA. In con-
trast, as noted above, GAs inhibit flowering.
Plant responses to GAs differ with species
140 and with the particular GA used; for exam-
25 50 75 100 ple, GA4+7 is more inhibitory to flowering of
Light transmitted by shades (%) apple than is GA3 or GA4 alone (Marino and
Fig. 7.3. Effects of shading on percentage of fruits Greene, 1981; Tromp, 1982), and some evi-
retained (top) and on mean fruit weight (bottom) dence suggests that other GAs can actually
(Jackson, 1975). promote flowering (Looney et al., 1985).
7.3 Flowering Habit, Flower Structure therefore contain up to ten seeds. Individual
carpels of some cultivars (e.g. ‘Northern Spy’)
Apple flowers can be initiated in both terminal may contain more than two ovules. Stamens
and axillary buds of both spurs and shoots. number approximately 20. The terminal, or
Cultivars differ in this respect, some flowering ‘king’, flower is the first to open and gives rise
primarily on terminal buds on shoots (e.g. to the largest fruits in ‘Delicious’ and some
‘Paulared’, ‘Rome Beauty’), some primarily on other cultivars. Flower clusters in lateral buds
terminal buds of spurs (e.g. ‘McIntosh’, open later than do those in terminal buds and
‘Delicious’). Some cultivars rarely or never generally produce smaller fruit.
flower on lateral buds (‘McIntosh’), while oth-
ers often do (‘Golden Delicious’, ‘Gala’). The
flower buds are mixed, containing both vege- 7.4 Pollination and Fertilization
tative and reproductive parts (Fig. 7.4). The
flower cluster is a cyme (terminal flower is the 7.4.1 Parthenocarpy
most advanced), is terminal within the bud
and may contain up to six individual flowers. Parthenocarpic cultivars of apple exist, but
Further vegetative growth develops from lat- they are of no economic value. The petals are
eral buds proximal to the flower cluster, lead- replaced by a second set of sepals and the
ing to formation of one or two bourse shoots. stamens by a set of ten carpels distal to the
The flower (Fig. 7.5) is epigynous, the normal five. Although pollination leads to
ovary being enclosed by non-ovarian tissue seed development, the flowers do not attract
(fused base of sepals, petals and stamens or bees and most of the fruits are seedless.
cortex of stem, depending on morphology Commercial cultivars are dependent upon
espoused) that remains attached to the ovary pollination. Although parthenocarpy can be
at harvest, giving rise to a ‘false’ fruit, or induced with growth regulators, primarily
pome (Fig. 7.6). A normal flower consists of GAs, response is limited and such treatments
five carpels, each with two ovules and five are not used commercially.
sepals, petals and styles. A normal fruit can
7.4.2 Pollination
Most apple cultivars require cross-pollina-

tion to set commercial crops of fruit (i.e. are
self-unfruitful) and, even in partially self-
fruitful cultivars, such as ‘Rome Beauty’,
‘Jonathan’, ‘Yellow Newtown’ and ‘York
Imperial’, cross-pollination is recommended.
Cross-incompatability (i.e. both cultivars
produce viable pollen, but neither will set
fruit when cross-pollinated) occurs in very
few combinations, such as ‘Early McIntosh’ ×
‘Cortland’. However, triploid cultivars are a
more common problem, as their pollen has
low viability. The haploid number in apple is
17; hence triploids have 51 chromosomes.
Therefore, the chromosomes are unequally
divided – or parts of them may occur in the
Fig. 7.4. Vertical section through an apple flower-
haploid cells – during meiosis. Such cultivars
bud (diagrammatic). Black appendages – bud (e.g. ‘Winesap’, ‘Mutsu’ (‘Crispin’),
scales; stippled – leaves; open – bracts and ‘Jonagold’) are ineffective as pollinizers for
subtended flowers (apical flower without a bract) other cultivars. Sports of cultivars, such as
(Abbott, 1970). the colour sports and spur strains of
158 F. Dennis, Jr
Dehisced Stigma
anther Style
Filament
Undehisced
anther
Nectar
Nectariferous droplet Petal
area Sepal
Ovary Ovule
Fig. 7.5. Structure of an apple flower (McGregor, 1976).
(a) (b)
Calyx lobe
Stamens
Styles
Endocarp
Seed
Cortex of
receptacle
Core line
(vascular
bundles)
Pith of
receptacle
Exocarp
and mesocarp
Fig. 7.6. Structure of a mature apple fruit. (a) Vertical section; (b) equatorial section (Robbins, 1933).
‘Delicious’, are not compatible (i.e. are self- Growers may wish to produce only one
unfruitful) with one another, as mutation cultivar in an orchard and use a minimum
does not affect the reproductive tissues. In number of trees as pollinizers. This can be
addition to being compatible, pollinizers done by planting pollinizers as every third
must bloom at the same time as the cultivar tree in every third row, providing a
being pollinated and should be annual, pollinizer adjacent to every tree of the main
rather than biennial, to ensure a supply of cultivar. To produce even fewer fruits of the
pollen each year. Nursery catalogues often pollinizer cultivar, grafts can be inserted on
contain compatibility charts, providing trees of the main cultivar at the same loca-
growers with information as to which culti- tions. Fruit of the cultivar used as the
vars are suitable as pollinizers. pollinizer should differ from those of the
main cultivar (e.g. yellow rather than red) to rent honey bees from apiculturists during
avoid mixing cultivars when harvesting. the bloom period, a minimum of four or five
Ornamental crab apples can be used as strong colonies per hectare being recom-
pollinizers in solid blocks of commercial cul- mended in mature orchards. The bees must
tivars, generally in orchards propagated on be removed from the orchard prior to appli-
dwarfing rootstocks, with trees planted in cation of insecticides. For maximum effec-
hedgerows. This avoids the need for having tiveness, the orchard floor should be mowed
more than one commercial cultivar in an to remove flowers (e.g. dandelion (Taraxacum
orchard and simplifies harvest and other officinalis L.)) that may compete with apple
practices. The crab apples are planted blossoms for the bees’ attention.
between trees of the commercial cultivar Poor set of ‘Delicious’ apple has been
within the rows and are pruned heavily so attributed to the presence of ‘basal gaps’
that they occupy little space. Three different between the filaments of the stamens,
cultivars are recommended, with bloom allowing bees to extract nectar without
times bracketing that of the cultivar being walking over the top of the flower
pollinated. They should be planted in each (Robinson, 1979). Evidence obtained by
row, as bees tend to travel up and down, DeGrandi-Hoffman et al. (1985), however,
rather than across, the rows, and should be did not support this conclusion.
offset in adjacent rows for optimum place- Timing is important in order for pollina-
ment. Blossom colour should be similar to tion to be effective. Williams (1966) proposed
that of the commercial cultivar, as individual the term ‘effective pollination period’ (EPP)
bees tend to work flowers of a single colour. for the interval during which pollination will
If pollen sources are lacking within the result in fertilization. Pollination must occur
orchard, several means are available for intro- within a given ‘window’ (the EPP) in order
ducing compatible pollen. Flowering branches for the pollen-tube to reach the ovule and for
of other cultivars (‘bouquets’) can be placed in fertilization to occur while the ovule is still
containers with the cut ends in water. receptive (Fig. 7.7). The length of the EPP
Alternatively, pollen can be purchased from varies with cultivar, tree condition and tem-
commercial companies and used in inserts perature. Ovule longevity is shorter follow-
placed at the entrance of honey bee (Apis mel-
lifera L.) hives. Bees exiting the hive unwit-
tingly pick up pollen and carry it to the Stigma receptive
flowers visited. Another device is available
that dusts the insects with pollen as they leave
the hive. Pollen can also be ‘dusted’ on trees
by dropping it into the draught created by an EPP
air-blast sprayer. Some growers use heli-
copters to apply pollen from the air, after mix-
ing it with a suitable diluent. However, the
‘target’ (stigmata of the flower) is very small; Ovule longevity
thus much pollen is wasted. As a last resort,
flowers can be pollinated by hand, but the
labour cost is high, even though only one or 0 2 4 6 8 10
two flowers in several clusters need be treated.
Days after anthesis
Apple pollen is heavy and is not carried
readily by the wind as is the pollen of some Fig. 7.7. The effective pollination period (EPP). The
ovule in this example is receptive for 8 days
tree species, such as conifers and nuts. The
following anthesis. Pollen-tube growth and
pollen is transferred primarily by insects,
fertilization require 2 days. Pollination at any time
especially honey and bumble (Bombus sp.) between 0 and 6 days will result in fertilization;
bees. During bloom, prolonged periods of thereafter the ovule is not receptive when the
cool weather or rain, which limit bee flight, pollen-tube reaches it. Thus, the EPP is 6 days
can be detrimental to fruit set. Fruit growers (= 8 2) (adapted from Williams, 1965).
160 F. Dennis, Jr
ing a heavy crop year, but may be prolonged (a)

in certain cases, at least, by application of
summer nitrogen (Williams, 1965). As the e
temperature following pollination rises, the e
pollen-tube grows more rapidly, within lim-
its, but the time during which the ovule is
receptive is reduced. Very high temperatures
n end in
are detrimental to fruit set. in n end
(b)
7.4.3 Fertilization n e
Once compatible pollen grains have been n

deposited on the stigma, they germinate and
the resulting pollen-tubes, each containing
three nuclei (tube nucleus and two genera- end
end
tive nuclei), grow down the style into the e
e
ovary. One tube enters the micropyle and
penetrates the ovule (Fig. 7.8), where it rup- a b c d
tures, releasing the two generative nuclei. Fig. 7.9. Stages in apple seed development.
One of these unites with the egg cell to pro- (a) Endosperm 24 (free-nuclear) and 31 days (cellular)
duce the diploid zygote and one unites with after fertilization. (b) Embryo and endosperm
the two polar nuclei in the embryo sac, pro- development at (a) 24, (b) 31, (c) 45 and (d) 76 days
ducing a triploid nucleus. The zygote after fertilization. in, integument; n, nucellus; end,
divides rapidly to produce the embryo, endosperm; e, embryo (Luckwill, 1959).
while the triploid nucleus divides to form a
free-nuclear, liquid endosperm (Fig. 7.9).
7.5 Fruit Development
Soon after fertilization occurs, the ovary and

surrounding receptacle tissues begin to grow
and the fruit has set. However, flowers in
Embryo sac
Antipodal cells
which fertilization did not occur soon fall
Ovary wall
and many developing fruits abscise before
Ovule
Polar reaching maturity. An abscission layer forms
nuclei Outer between the base of the pedicel and the clus-
integument
Inner ter base. Most abscission occurs within the
integument first 4–6 weeks of growth, culminating in the
Nucellus ‘June’ drop. ‘King’ flowers set better than do
Exine
lateral ones, and those in terminal clusters
Egg (outside) better than those in lateral clusters. Cultivars
cell Synergid 2X differ considerably in set; ‘Golden Delicious’,
cells
Funiculus ‘Paulared’ and ‘Fuji’ set a higher proportion
Intine of flowers than do ‘McIntosh’ and
Micropyle
(inside) ‘Delicious’. For this reason, the former tend
1X
to be biennial, the latter annual, bearers.
Tube nucleus
Slow-growing fruits and/or those with few
seeds generally drop first, as they are less
Sperm nuclei competitive. When initial set is light, subse-
quent abscission is less intense, primarily
Fig. 7.8. The process of fertilization (diagrammatic) because there is less competition among
(Dennis, 1996). fruitlets. Cloudy weather limits photosyn-
thesis, thus favouring abscission, and fruit results in maximum economic value of the
set is generally greater in western regions of crop as a whole. When set is below the opti-
the USA where sunlight is abundant than in mum, fruit size is large and therefore the
the humid and cloudy east, and in well- individual fruits are more valuable, but this
exposed portions of the tree than in the usually does not compensate for the smaller
shaded, inner portions (Fig. 7.3). Negative number of fruits. Early thinning can be
correlations between temperature prior to accomplished either manually, which is
bloom and fruit set have been reported for expensive, or by using chemicals that induce
several cultivars and locations (Jackson and abscission of a portion of the flowers or fruits
Hamer, 1980). (see below and Chapter 16). Growers often
Several cultural practices can be used to thin with specific chemicals and then hand-
improve set. Pruning opens the tree to light, thin to remove excess fruits. Early thinning
thus avoiding excessive shading. This is less also increases flower initiation for the next
of a problem with trees propagated on year’s crop, thereby reducing biennial bear-
dwarfing rootstocks than with large trees on ing. Thinning also reduces the danger of
seedling rootstocks. The rootstock can also limb breakage from excessive weight on
have a further major effect, with set being limbs, and can improve fruit colour and
greater on less vigorous stocks. The addition quality by reducing reciprocal shading
of major elements, particularly nitrogen, is among fruits in the same cluster and by
critical for good fruit set in most orchards. increasing the fruit’s access to carbohydrates,
Nitrogen should be applied at least 1 month which are required for both growth and
before flowering so that it will be available at anthocyanin formation.
the critical time. Sprays of urea can improve
fruit set once sufficient leaf surface is avail-
able to absorb this source of nitrogen; how- 7.6 Seed and Fruit Growth
ever, results have been variable. Although
boron sprays improve set in some plum About 4–6 weeks after fertilization, the
(Prunus domestica L.) cultivars, apple is not endosperm becomes cellular and soon fills
responsive. Scoring or ringing the bark of the much of the developing ovule (seed) as it
trunk or of large scaffold limbs can reduce grows at the expense of the nucellus (Fig.
the June drop, thereby increasing final set. 7.9). The embryo develops more slowly, but
This is usually done with young trees, in gradually consumes the endosperm and
which fruit set can be poor. The treatment is occupies most of the seed at maturity.
effective only within a few weeks of bloom During the first 3–4 weeks of growth, both
(see also Chapter 14). cell division and cell expansion are occurring
Increasing fruit set when conditions for in the fruitlet. Thereafter, increase in size is
pollination are poor is more difficult than is almost entirely the result of expansion of cells
reducing crop load when it is excessive. and intercellular spaces, except in the epider-
Although GAs can improve fruit set in some mis, where cell division continues. Although
species, they do not have this effect in apple. cell expansion contributes much of the vol-
One of the most effective chemicals tested to ume, cell division is critical in determining
date for increasing set is aminoethoxyvinyl- final size. Warm temperatures early in the
glycine (AVG), an inhibitor of ethylene season stimulate fruit growth and increase
biosynthesis. Although this compound, mar- ultimate size (Warrington et al., 1999), but
keted as Retain®, is currently used commer- shorten the period of cell division. The
cially for delaying maturity and reducing carpellary (ovary) tissue stops growing
preharvest abscission of apple fruits (see approximately 6 weeks after bloom, whereas
Chapter 17), it is not used commercially to the cortex (fused base) continues to expand
improve fruit set. (Fig. 7.10). Fruit growth continues as fruit
When fruit set is excessive, the fruit must mature; delaying harvest increases fruit size,
be thinned to allow more of them to grow to but over-maturity and excessive preharvest
marketable size, as an optimum crop load drop may occur.
162 F. Dennis, Jr
example, flowers initiated during a heavy

crop year may have fewer cells than do those
) (mm)
Diameter
60 initiated during a light crop year. Large fruits
Entire fruit
contain more cells than do small ones, and
50 early hand-thinning increases fruit size pri-
Length
) or diameter (
marily by increasing cell number. Fruit

40 growth can be limited by water and nutrient
supply and by any factors that affect photo-
30 synthesis, such as low temperature, cloudy
weather and insect injury to the leaves.
20 Carpel Fruit size at harvest can be predicted by
Length (
measuring fruit diameter during the grow-

10 Nucellus / integuments ing season and projecting the growth curve
to harvest. This procedure is useful in evalu-
Embryo
0 ating how best to use the fruit (fresh fruit,
May June July Aug. Sept. processing, etc.) and can be helpful in decid-
Time of sampling ing when and how much to thin (Fig. 7.11).
Fig. 7.10. Growth of seed and fruit tissues in a Fruit shape varies with cultivar and stage
‘McIntosh’ apple at Geneva, New York State, from of development and ranges from oblate (flat)
fertilization until maturity (Tukey and Young, 1942). to round to conic. The ratio of length to
diameter (L/D ratio) declines as fruits
enlarge, resulting in lower ratios for larger
The major factors determining ultimate fruits (Fig. 7.12). Thus heavy cropping leads
size are cultivar and crop load; exposure to to a high L/D ratio. In ‘Delicious’, cool tem-
light also plays an important part in deter- peratures during the first few weeks after
mining final size (Fig. 7.4; see also Chapter bloom stimulate growth of the apical portion
9). Seed number, rootstock, fruit position in of the fruit, leading to a high L/D ratio; in
the cluster and on the shoot and spur size warmer areas the fruit are often nearly round
also affect the rate of fruit enlargement. Fruit (Fig. 7.13). The ‘typey’ (high L/D ratio)
size has been positively correlated with seed appearance of Washington State ‘Delicious’
number in some cultivars. Size is greater on is a selling-point for such fruit and their total
certain rootstocks than on others, possibly weight is greater than non-typey fruit of the
because branch angle is affected, which in same diameter. Rootstock and position on
turn influences light penetration. In the cluster can also affect the ratio; L/D
‘Delicious’, the ‘king’ flower produces a ratios of ‘king’ fruit are smaller than those of
noticeably larger fruit than do the lateral lateral ones, at least in ‘Delicious’.
flowers, primarily because of larger cells, Promalin®, a mixture of benzyladenine (BA)
whereas in ‘McIntosh’ and ‘Empire’ the dif- and GA4+7, is used commercially to stimulate
ference is much less pronounced. When only growth of the apical portion of the fruit to
one ‘Delicious’ fruit is left per cluster, size produce a ‘typey’ appearance.
differences between lateral and terminal
fruits are negligible; however, if both a termi-
nal and a lateral fruit are allowed to develop 7.7 Fruit Maturation
on the same cluster, growth of the latter is
retarded, indicating the greater ability of the Cultivars differ widely in time of ripening;
terminal fruit to attract nutrients (Black and some (e.g. ‘Lodi’) ripen within 60 days after
Bukovac, 1996). Size increases with spur leaf full bloom (DAFB) while others require 180
area; fruits developing from lateral flower- days or more to mature (e.g. ‘Granny
buds are smaller than those produced from Smith’). Rootstock also can affect time to
terminal ones, probably because leaf area is maturity, but has much less effect than does
smaller, although the anatomy of the cluster cultivar. Heavy cropping delays maturation.
base and flower may be involved as well. For Climatic factors also affect maturation, tem-
100
Relative benefit from thinning (%)

80
Winesap
60
40
Delicious
20
0
0 20 40 60 80 100 120 140 160 180
Days after full bloom
Fig. 7.11. Calculated percentage benefit, in terms of fruit size, of thinning apples at various times after full
bloom (Batjer et al., 1957).
1.7 Several methods are available to deter-

1.6 mine and/or predict optimum time of har-
vest. These include methods based on
1.5 temperatures early in the season, the ethyl-
Length/diameter ratio
1.4 ene content of/production by the fruit and

1.3 Pear other properties, such as firmness and con-
centration of soluble solids.
1.2 Chemicals can be used to either hasten or
Peach delay maturation (see also Chapter 17).
1.1
Apple Apple is a climacteric fruit – a burst of car-
1.0
bon dioxide and ethylene production,
0.9 termed the ‘climacteric’, occurs in the fruits
as they ripen – and ethylene acts as a ripen-
10
30 50 70 90 110 130 150 ing hormone. Sprays of ethephon, which
Days after full bloom releases ethylene within the treated tissues,
Fig. 7.12. Change in L/D ratio of apple, peach and can be used to hasten maturity in order to
pear fruits during fruit development (Westwood, obtain higher prices. Ethephon is generally
1993). used only with the earliest cultivars.
Because it stimulates ripening, the fruits
cannot be stored for long periods, but such
perature and solar radiation being the most early cultivars generally do not have a long
important. Low spring and summer temper- storage life, even without ethephon treat-
atures delay maturation; the time from full ment. Furthermore, the prices for them fall
bloom to harvest of a given cultivar is quickly as better, later-maturing, cultivars
longer in Norway than in Italy because of become available.
the cooler climate. Exposure to sunlight Chemicals that delay maturation – and
not only increases red coloration; it also retard preharvest fruit abscission – include
increases fruit sugar content because of its Alar®, which, as noted above, is no longer
effects in stimulating photosynthesis in adja- available for commercial use, and Retain®
cent leaves. Fruits in heavily shaded parts of (= AVG, mentioned above). To be effective,
the tree are not only smaller and greener but these chemicals must be applied approxi-
less mature as well. mately 1 month before harvest. Delaying
164 F. Dennis, Jr
1.00
Fruit shape (length/diameter ratio)

0.96
0.92
0.88
0.84
0.80
1667 2167 2667
Degree-days above 5C
Fig. 7.13. Effect of temperature during the growing season on final length/diameter ratio of ‘Delicious’
apple fruit (Westwood, 1993).
maturity prolongs both the harvest season and inhibition of subsequent flowering –
and the fruit’s storage life and allows more hence biennial bearing – unless the crop is
time on the tree for the fruit to develop both reduced by early thinning. Pollination and
size and red colour. Certain auxins used to fertilization are essential for fruit set.
delay abscission, such as naphthaleneacetic Although some cultivars are self-fruitful,
acid (NAA) and 2,4,5-trichlorophenoxypro- cross-pollination is required in most and is
pionic acid (2,4,5-TP), both delay abscission usually advantageous even in self-fruitful
and hasten maturity. However, the former ones. Bees are the primary pollinators, and
effect is short-lived; if the fruits are not har- climatic conditions during bloom are critical
vested within 10–14 days of treatment, abcis- for fruit set. When fruit set is excessive, the
sion can be stimulated rather than inhibited. fruits must be thinned mechanically or
chemically to encourage fruit growth and
flower-bud formation. Cultivars differ in the
7.8 Summary time required for maturation, some ripening
in midsummer and some in late autumn.
In crops whose marketable organs are fruits Preharvest drop can be a problem in some
and/or seeds, such as apple, flowering, fruit years and with some cultivars. Chemicals
set and development are critical for eco- can be used either to hasten ripening or to
nomic return. Some apple cultivars tend to delay it, allowing growers to harvest earlier
set too many fruits resulting in small size or later than would otherwise be the case.
Further Reading
Buban, T. and Faust, M. (1982) Flower bud induction in apple trees: internal control and differentiation.
Childers, N.F., Morris, J.R. and Sibbett, G.S. (1995) Modern Fruit Science: Orchard and Small Fruit Culture,
10th edn. Horticultural Publications, Gainesville, Florida, pp. 92–105.
Dennis, F.G., Jr (1979) Factors affecting fruit set in apple, with emphasis on Delicious. Horticultural
Reviews 1, 385–422.
Dennis, F.G., Jr (1985) Apple. In: Monselise, S.P. (ed.) Handbook of Fruit Set and Development. CRC Press,
Boca Raton, Florida, pp. 1–44.
Dennis, F.G., Jr (1996) Fruit development. In: Maib, K. (ed.) Tree Fruit Physiology: Growth and Development.
Washington State Fruit Commission, Yakima, Washington, pp. 109–116.
Feucht, W. (1976) Fruitfulness in Pome and Stone Fruits. Extension Bulletin 665, Washington State
University, 32 pp. (translated from German).
Forshey, C.G. and Elfving, D.C. (1989) The relationship between vegetative growth and fruiting in apple
trees. Horticultural Reviews 11, 229–287.
Greene, D.W. (1966) Flower development. In: Maib, K. (ed.) Tree Fruit Physiology: Growth and Development.
Nyeki, J. and Soltesz, M. (1996) Floral Biology of Temperate Zone Fruit Trees and Small Fruits. Akademiai
Kiado, Budapest, 377 pp.
Pratt, C. (1988) Apple flower and fruit: morphology and anatomy. Horticultural Reviews 10, 273–308.
Williams, R.R. (1970) Factors affecting pollination in fruit trees. In: Luckwill, L.C. and Cutting, C.V. (eds)
Physiology of Tree Crops. Academic Press, London, pp. 193–207.
References
Abbott, D.L. (1970) The role of budscales in the morphogenesis of the apple fruit bud. In: Luckwill, L.C.
and Cutting, C.V. (eds) Physiology of Tree Crops. Academic Press, New York, pp. 65–82.
Abbott, D.L. (1977) Report of the Long Ashton Research Station for 1976. Long Ashton, Bristol, UK,
pp. 162–176.
Batjer, L.P., Billingsley, H.D., Westwood, M.N. and Rogers, B.L. (1957) Predicting harvest size of apples at
different times during the growing season. Proceedings of the American Society for Horticultural Science
70, 46–57.
Black, B. and Bukovac, M.J. (1996) Plant growth regulator application technology, uptake and action. In:
Maib, K. (ed.) Tree Fruit Physiology: Growth and Development. Washington State Fruit Commission,
Yakima, Washington, pp. 41–50.
Chan, B.G. and Cain, J.C. (1967). The effect of seed formation on subsequent flowering in apple.
Proceedings of the American Society for Horticultural Science 91, 63–68.
DeGrandi-Hoffman, G., Hoopingarner, R. and Baker, K.K. (1985) The influence of honey bee ‘sidework-
ing’ behavior on cross-pollination and fruit set in apples. HortScience 20, 397–399.
Dennis, F.G., Jr (1996) Fruit set. In: Maib, K. (ed.) Tree Fruit Physiology: Growth and Development.
Dennis, F.G., Jr and Neilsen, J.C. (1999) Physiological factors affecting biennial bearing in tree fruit: the
role of seeds in apple. HortTechnology 9, 317–322.
Edwards, G.R. and Notodimedjo, S. (1987) Defoliation, bending, and tip pruning of apple under tropical
conditions. Acta Horticulturae 199, 125–127.
Erez, A., Fishman, S., Gat, Z. and Couvillon, G.A. (1988) Evaluation of winter climate for breaking bud
rest using the dynamic model. Acta Horticulturae 232, 76–89.
Harley, C.P., Magness, J.R., Masure, M.P., Fletcher, L.A. and Degman, E.S. (1942) Investigations on the
Cause and Control of Biennial Bearing of Apple Trees. Technical Bulletin 792, United States Department
of Agriculture, Washington, DC.
Hill-Cottingham, D.G. and Williams, R.R. (1967) Effect of time of application of fertilizer nitrogen on the
growth, flower development and fruit set of maiden apple trees, var. Lord Lambourne, and on the
concentration of total nitrogen within the tree. Journal of Horticultural Science 42, 319–338.
Jackson, J.E. (1975) Effects of light intensity on growth, cropping, and fruit quality. In: Pereira, H.C. (ed.)
Climate and the Orchard: Effects of Climatic Factors on Fruit Tree Growth and Cropping in South-eastern
England. Research Review No. 5, Commonwealth Bureau of Horticulture and Plantation Crops, East
Malling, Maidstone, UK, pp. 17–31.
Jackson, J.E. and Hamer, P.J.C. (1980) The causes of year-to-year variation in the average yield of Cox’s
Orange Pippin apple in England. Journal of Horticultural Science 55, 149–156.
Looney, N.E., Pharis, R.P. and Noma, M. (1985) Promotion of flowering in apple trees with gibberellin A4
and C-3 epi-gibberellin A4. Planta 165, 292–294.
Luckwill, L.C. (1959) Fruit growth in relation to internal and external chemical stimuli. In: Rudnick, D.
(ed.) Cell, Organism and Milieu. Ronald Press, New York, pp. 223–251.
McGregor, S.E. (1976) Pollination of economic crops. Apple. In: Handbook 496. United States Department
of Agriculture, Washington, DC, pp. 81–88.
166 F. Dennis, Jr
Marino, F. and Greene, D.W. (1981) Involvement of gibberellins in the biennial bearing of ‘Early
McIntosh’ apples. Journal of the American Society for Horticultural Science 106, 593–596.
Neilsen, J.C. and Dennis, F.G., Jr (2000) Effects of seed number, fruit removal, bourse shoot length and
crop density on flowering in ‘Spencer Seedless’ apple. Acta Horticulturae 527, 137–146.
Richardson, E.A., Seeley, S.D. and Walker, D.R. (1974) A model for estimating the completion of rest of
‘Redhaven’ and ‘Elberta’ peach trees. HortScience 9, 331–332.
Robbins, W.W. (1933) The Botany of Crop Plants, 3rd edn. P. Blakiston’s Son and Co., Philadelphia,
Pennsylvania, 674 pp.
Robinson, W.S. (1979) Effect of apple cultivar on foraging behavior and pollen transfer by honey bees.
Tromp, J. (1976) Flower-bud formation and shoot growth in apple as affected by temperature. Scientia
Tromp, J. (1982) Flower bud formation in apple as affected by various gibberellins. Journal of Horticultural
Science 57, 277–282.
Tukey, H.B. and Young, J.O. (1942) Gross morphology and histology of the developing fruit of the apple.
Botanical Gazette 104, 1–25.
Warrington, I.J., Fulton, T.A., Halligan, E.A. and de Silva, H.N. (1999) Apple fruit growth and maturity
are affected by early season temperatures. Journal of the American Society for Horticultural Science 124,
468–477.
Weinbaum, S.A., DeJong, T.M. and Maki, J. (2001) Reassessment of seed influence on return bloom and
fruit growth of ‘Bartlett’ pear. HortScience 36, 295–297.
Weinberger, J.H. (1950) Chilling requirements of peach varieties. Proceedings of the American Society for
Westwood, M.N. (1993) Temperate-zone Pomology: Physiology and Culture, 3rd edn. Timber Press, Portland,
Oregon, 523 pp.
Williams, R.R. (1965) The effect of summer nitrogen applications on the quality of apple blossom. Journal
of Horticultural Science 40, 31–41.
Williams, R.R. (1966) The effective pollination period for some apple and pear varieties. In: Report of the
Long Ashton Research Station for 1965. Long Ashton, Bristol, UK, pp. 136–138.
8 Water Relations of Apples
Alan N. Lakso
Fruit Crop Physiology Program, Cornell University, Department of Horticultural
Sciences, New York State Agricultural Experiment Station, Geneva, New York State,
USA
8.1 Basics of Water Relations of Apple Trees 168

8.2 Concepts of Water Relations 168
8.2.1 Water-potential components 168
8.2.2 Plant hydraulic resistance 169
8.3 Measurement of Apple-tree Water Status 170
8.3.1 Water-potential measurements 170
8.3.2 Trunk- and fruit-diameter monitoring 171
8.3.3 Stable-isotope discrimination 171
8.4 Root-system Characteristics and Influences on Water Relations 171
8.4.1 Availability of soil water and nutrients 172
8.4.2 Implications of plant resistance and water status 172
8.4.3 Root growth and water uptake 172
8.4.4 Mycorrhizae and water uptake 173
8.5 Water Use by Apple Orchards 173
8.6 Factors that Affect Water Use in Apple Orchards 173
8.6.1 Energy, radiation and humidity 173
8.6.2 Leaf area 174
8.6.3 Crop structure and boundary layers 174
8.6.4 Factors affecting stomatal opening 175
8.6.5 Canopy form, spacing and light interception 175
8.6.6 Interactions with drought 175
8.7 Water Deficits and Apple-tree Growth, Cropping and Physiology 175
8.7.1 Timing of stress 176
8.7.2 Vegetative growth 176
8.7.3 Fruit growth, quality and postharvest effects 177
8.7.4 Gas exchange 178
8.8 Integration of Water-stress Effects 179
8.8.1 Drought avoidance or escape 179
8.8.2 Tolerance by maintaining high water potentials 179
8.8.3 Tolerance of low water potentials 179
8.9 Effects of Rootstocks on Apple Water Relations and Drought Tolerance 180
8.9.1 Controlled drought-response studies 180
8.9.2 Water-relations studies in the field 180
8.9.3 Indirect effects 181

168 A.N. Lakso
8.10 Management of Water Relations in the Field 181

8.10.1 Orchard water balance 181
8.10.2 Irrigation management 182
8.10.3 Plant-based methods for irrigation scheduling 184
8.10.4 Deficit irrigation with apples 185
8.11 Needs for Integrative Approach to Water Relations 186
8.11.1 Modelling water relations 186
8.12 Conclusions 187
8.1 Basics of Water Relations of Apple whether they increase or decrease water
Trees activity:
Total = Osmotic + Turgor + Matric +
Water relations are very important to the func-
() () (+) ()
tion of the apple tree, as water is the greatest
Gravitational
component of the tree by mass and almost all
critical processes can be limited by inappropri- ()
ate water status. The essential role of water, Normally, gravitational potential changes
however, does not mean that the water is only by 0.01 MPa m−1 above the ground, so it
always limiting or is regulating variations in can be ignored except in very tall trees.
productivity. This chapter will address some Similarly, the matric potential is the reduction
of the times and conditions (natural and cul- in potential due to interactions of water with
turally imposed) under which the water rela- surfaces, so it is important in soils. However,
tions may be controlling tree behaviour. it is usually included in the osmotic-potential
Specific reviews are available on water rela- measurement for plant cells.
tions and irrigation of fruit-trees (Elfving,
1982; Jones et al., 1985; Bravdo and Proebsting,
1993) and on apple (Landsberg and Jones, 8.2.1.1 Total water potential
1981; Lakso, 1994; Behboudian and Mills,
Total water potentials (w) are controlled by
1997), so this chapter will focus on an interpre-
the balance of osmotic and turgor potentials.
tative review of more recent information.
Within the tree, total water potentials
describe the gradients for water movement,
8.2 Concepts of Water Relations with water moving from high (less negative)
to low water potentials. So the primary
Water relations of plants are typically importance of total water potentials is to
described in terms of the thermodynamics of determine the direction of water movement
water activity and are expressed in terms of and strength of gradient for that movement.
megapascals (1 MPa = 10 bars as used in
older papers). The components of water rela- 8.2.1.2 Osmotic potential
tions and measurement methods will be
reviewed briefly to provide a basis for later Osmotic potential or solute potential (s) is
discussions. The reader is referred to excel- the lowering of water activity by the interac-
lent books by Nobel (1991) and Jones (1992) tion of water with solutes in the cell. It is a
for more detailed discussions on water rela- ‘colligative’ property, which means that it
tions, but a brief review will follow. depends on the concentration of the solutes,
not the size of the molecules. So a mole of
potassium ions has the same effect on
8.2.1 Water potential components osmotic potentials as a mole of glucose.
Adjustments in the osmotic potential of a cell
The total water potential is made up of sev- or tissue modify the relationship between
eral components of varying sign as to total and turgor potentials. At a constant
Water Relations of Apples 169
total water potential, a more negative plant resistance is that it affects the relation-
osmotic potential due to accumulation of ship between the water-potential gradient
solutes will increase turgor. For example, the and the transpiration flux of water (E). This
variation in turgor and opening of stomata is relationship is expressed as an analogy to the
controlled primarily by fluxes of potassium electrical Ohm’s law for the relationships
ions into and out of the guard cells. among a potential gradient (voltage), a resis-
Conversely, as water potential becomes more tance and a flux (current):
negative with drought stress, leaves of apple
E = d w / R
can lower the s (i.e. it becomes more nega-
tive) by accumulating sugars and other Compared with many plants, such as
solutes to maintain turgor and leaf function, annuals, apple trees have a high hydraulic
as will be discussed later (Goode and Higgs, resistance (Fig. 8.1). This means that a rela-
1973; Lakso et al., 1984). Apple fruits also tively large gradient of w is needed to
accumulate many solutes during develop- move enough water through the tree to
ment that affect the fruit s and fruit water maintain any given transpiration rate. The
relations. Additionally, the hydrolysis of intercept at E = 0 is controlled by the soil
starch to sugars as the fruit matures lowers water potential, the minimum value at mid-
the s (i.e. it becomes more negative) with- day is determined by E, while the slope of
out requiring imported carbohydrates. the relationship is the tree resistance, which
is mostly in the apple root system
(Landsberg and Jones, 1981).
8.2.1.3 Turgor potential
Consequently, for similar transpiration
Turgor or pressure potential (p) refers to the rates, apple trees, and many other trees, will
changes in water activity due to the positive show a much greater diurnal decrease in leaf
pressures that develop as water is drawn into water potential than do annuals. This is seen
cells by the osmotic reduction of water in the typical strong diurnal decline of leaf
activity. Turgor pressure is critical as the water potential of apple leaves on a sunny
energy source for expansive growth of cells day (Fig. 8.2). Because the predawn water
and for tissue turgidity of all parts of the tree. potential is determined by the soil water
Many plant processes seem to sense turgor, potential if the E rate is near zero, predawn
although the mechanisms of sensing are not leaf water potentials are used as estimates of
well known. p normally changes with total
water potential unless there is a compen-
0
satory change in osmotic potential that may Annual
help maintain p at a more constant level.
Leaf water potential (MPa)
–1 Apple
8.2.1.4 Relative water content wet
soil
Relative water content refers to the amount
of water that a cell or tissue holds as a per-
centage of what it could hold when fully –2
hydrated (analogous to relative humidity). It
is related to the total and turgor potentials, Apple
dry
but not directly. soil
–3
0 1
Relative transpiration rate
8.2.2 Plant hydraulic resistance
Fig. 8.1. General relationship of leaf total water
Plant hydraulic resistance (R) refers to the potential of the top of the tree to the transpiration
resistance within the vascular system to rate. The steeper the slope, the greater the hydraulic
water movement along the gradients of total resistance. Soil water potential affects the intercept
water potential. The main significance of where E = 0.
170 A.N. Lakso
Water potential (MPa) –1.0
–2.0
6 8 10 12 14 16 18 20
Time (h)
Fig. 8.2. Diurnal patterns of apple leaf total water potentials on trees irrigated to the soil (triangles) or by
misting the canopy (solid circles) compared with unirrigated controls (open circles) (adapted from Brough et
al., 1986, with permission).
effective soil water potential around the 8.2.2.1 Water-use efficiency (WUE)
roots. This can be useful, as representative
The WUE of a plant is typically the carbon
placement of soil moisture measuring
gained per unit of water lost. It may be
devices is difficult since apple root systems
expressed in instantaneous net photosynthe-
have very low density and are erratic in dis-
sis rate per transpiration rate, or it may be a
tribution (Atkinson, 1980; Hughes and
long-term integral of dry matter per volume
Gandar, 1993; de Silva et al., 1999).
of water loss. In apple trees in the field, we
Since E varies with the evaporative condi-
have found good WUE, since the stomata
tions, the leaf w also depends strongly on
maintain an optimal conductance that is
evaporative conditions of the atmosphere, as
tightly coupled to the photosynthesis rate
shown by the effectiveness of aerial misting
(Lakso, 1994). The net effect is that the stom-
on reducing the diurnal pattern of leaf water
atal conductance adjusts so that, for exam-
potential (Goode et al., 1979; Brough et al.,
ple, reductions in the photosynthesis rate
1986; Fig. 8.2). This is not the case for a plant
due to lack of crop or some form of girdling
that has low hydraulic resistance, such as the
are matched by reduction in conductance.
sunflower (Fig. 8.1). In apple, however,
atmospheric conditions are much more
important to the control of w (see Jones et
8.3 Measurement of Apple-tree Water
al., 1985, for a more detailed discussion). This
Status
means that:
• Soil moisture measurements cannot be 8.3.1 Water-potential measurements
used alone to estimate midday water
potentials, which depend more on evapo- The most common method of measuring tree
rative demands. water status has been to estimate exposed-leaf
• Water potentials will be highly variable if total water potential (w) with a Scholander
the conditions are variable. pressure chamber (also known as the pressure
• Therefore, the uniformity of environ- bomb). Although many leaf processes, such as
mental conditions is critical for measure- stomatal opening and photosynthesis, are cor-
ments when comparing the water related with w, the limitations of using w
potential effects of treatments (i.e. mea- alone include: (i) significant osmotic adjust-
surements must be compared under sim- ment in the apple, which can change critical
ilar conditions). levels of w (Lakso et al., 1984); (ii) variability
• Cool, humid conditions will ameliorate a due to individual leaf exposure and transpira-
soil drought while hot dry conditions will tion rates so that exposed-leaf w may not
aggravate the drought effects (Sritharan represent shaded leaves, fruit or shoot tips,
and Lenz, 1989). which do not transpire as much (Higgs and
Jones, 1990); and (iii) stomatal closure may monitoring these changes and proposed this
reduce transpiration enough to stabilize as a method for irrigation timing. Since then
exposed-leaf w so that w is not related to several studies have evaluated this method
internal water status (Jones et al., 1983). (Powell, 1976a; Assaf et al., 1982; Higgs and
Another common method, predawn leaf Jones, 1984; Iancu, 1985), although apparent
w, indicates effective potential at the contradictions in behaviour need to be recon-
soil/root interface under near-zero transpira- ciled (Huguet et al., 1992; Bonany et al., 2000)
tion if the soil moisture is homogeneous. before this method can be used in the field.
With deep root systems, especially with drip
irrigation, the predawn reading represents
the wettest part of the soil, not the average 8.3.3 Stable isotope discrimination
(Jones, 1990; Ameglio et al., 1999).
Consequently, the predawn value may not be This method is based on the discrimination
a good measure of midday stress under high against stable isotopes of different molecular
transpiration flow if only a small portion of weight (13C, 16O and 2H) during diffusion
the soil is wet. Consequently, a more integra- and exchange processes in the soil and in the
tive method is the estimation of midday stem plant (see Ehleringer et al., 1993).
potential (stem) by enclosing a leaf in a plas- Discrimination of 12C and the heavier 13CO2
tic bag with foil to shade the bag (Powell, during gas exchange is an integrator of vari-
1974; Olien and Lakso, 1986; Jones, 1990; ations in water-use efficiency, a major advan-
McCutchan and Shackel, 1992). This stops tage over the instantaneous methods
transpiration and allows the leaf to equili- generally used. In apple, Behboudian’s group
brate with the water potential in the stem at (Behboudian et al., 1994; Mills et al., 1998)
that point. The stem is a more integrative found that high root temperatures reduced
and stable measurement since it is influenced transpiration and discriminated against 13C
by all the leaves and organs of the branch. It and that deficit irrigation had similar effects,
is also a better estimate of the potential expe- although the differences were quite small.
rienced by fruits, shoot tips and other organs The need for an integrator of stress is even
that do not transpire rapidly. The stem value greater with apples, since the water relations
is probably the best single measure of plant of apple trees are so dependent on the
water status and is recommended (Naor, dynamic evaporative conditions; instanta-
2000). A promising approach is to integrate neous measurements during stable midday
midday stem potentials over a season into a conditions may not represent a large propor-
‘crop water-deficit index’, which was found tion of the long-term conditions. However, it
to be well correlated with apple fruit growth is possible that the carbon isotope method
and weight at harvest (Ebel et al., 2001). may not be as promising as hoped because in
the field apple leaf stomata are well coupled
to photosynthesis and mature leaves can
8.3.2 Trunk- and fruit-diameter monitoring osmotically adjust (Lakso, 1994).
The shrinking and swelling of apple-tree

trunks and fruit in relation to soil moisture 8.4 Root-system Characteristics and
deficits and evaporative demand have been Influences on Water Relations
recognized for many years (Furr and Magness,
1930; Verner, 1937; Harley and Masure, 1938; Two of the key characteristics of the apple tree
Taerum, 1964). The advantages are: (i) the root system of relevance to water relations
measurement can be monitored continuously are: (i) an extremely low root-length density
so is less dependent on the environment at in soil; and (ii) a very non-uniform root distri-
any one time; and (ii) the trunk and fruit better bution. Several studies of apple root systems
integrate the whole-tree water status than any have shown that apple root-length densities
single leaves. Almost 40 years ago Tukey are very low compared with grasses
(1963) developed an early electronic device for (Landsberg and Jones, 1981; Hughes and
172 A.N. Lakso
Gandar, 1993; de Silva et al., 1999; Green and face of the root and soil). Soil resistance for
Clothier, 1999). There are several implications water movement increases markedly as the
of such low root density and erratic rooting. soil dries and as the distance of travel to the
root increases. Based on estimates of crop
water use in arid climates, apple orchards in
8.4.1 Availability of soil water and nutrients mid-season transpire almost as much as grass
fields (Ley, 1994b; Allen et al., 1998; Hanson et
One important implication is that the effec- al., 1999), even though the root density of the
tive soil volume that is explored for water grass may be 1000 times higher than that of
and nutrients is reduced, especially in rela- the apple (Landsberg and Jones, 1981).
tion to non-mobile nutrients. There may be Atkinson and Wilson (1980) proposed that
relatively large portions of the potential soil the very low root-length density, combined
volume that are not explored by fine apple with the high water uptake required of each
roots. This can be seen in the results of root, will lead to localized drying in the rhi-
Atkinson and Wilson (1980), who found a rel- zosphere during times of high transpiration
atively high percentage of soil cores under at midday. This would occur if the soil could
mature apple trees at different distances and not supply water to the rhizosphere as fast as
depths to have no or few apple roots in them. it was taken up by the roots. Landsberg and
The sparse rooting also means that the Jones (1981) calculated that the soil–plant
average distance that is required for water resistance would increase much more rapidly
and nutrients to travel through the soil is in drying soil for plants with low root densi-
much greater for apple root systems than for ties such as apples. Direct evidence is lacking,
dense uniform root systems, such as grass. In but Bonany and Camps (1998) found that tree
competitive interactions of apple roots with growth and fruit size increased with irriga-
weeds or cover crops, the apple roots are tion rates up to 150% of crop evapotranspira-
generally very poor competitors. Normally, tion (ET). This has also been seen in almonds,
apple roots will not be able to establish in grapes and citrus (Hutmacher et al., 1994;
zones of weed or cover-crop root growth and Williams, 1996; Parsons et al., 2001). This
generally will require at least a minimum question deserves more attention to deter-
amount of surface soil with no competing mine its importance and if there are ways to
plants even with irrigation (Merwin and Ray, manage the effect for benefit.
1997). This may further reduce the available
soil volume for water and nutrient uptake.
However, it may also provide a management 8.4.3 Root growth and water uptake
tool to purposely restrict tree growth or, con-
versely, to improve growth by removing the In most studies of water and nutrient uptake,
competition of plants between the rows. In it is concluded that young, white roots are the
arid climates in which the tree depends on most efficient, especially for phosphorus
irrigation for most of the season, root distrib- uptake, although older roots are still quite
utions may concentrate in the wetted zone, active (Atkinson and Wilson, 1980). This
especially if nutrients are supplied by ferti- implies that flushes of new root growth
gation (Huguet, 1976; Levin et al., 1979, 1980; should stimulate better water and nutrient
Bravdo et al., 1992; Neilsen et al., 2000). uptake. Surprisingly, there is little direct evi-
dence in apple for this conclusion. In part this
is due to little knowledge of when roots in fact
8.4.2 Implications for plant resistance and are growing during the season. The many
water status detailed studies of apple root growth at the
East Malling Research Station rhizotron over
As discussed above, the apple root system several decades (see Atkinson, 1980, 1983) pro-
has a quite high hydraulic resistance. An vided much information, but were limited to
additional component of this can be the resis- one site. Recent studies in New York State
tance of the soil in the rhizosphere (the inter- found that, over several seasons, new root
production generally did not occur until about 8.5 Water Use by Apple Orchards
1 month after bloom and the great majority of
growth was completed in 60–80 days (Psarras The most direct method to measure water
et al., 2000; A. Lakso and K.-T. Li, unpublished use is by lysimeter, which can monitor either
data). However, in a warm dry year with weight loss (weighing lysimeter) or the
heavy crop loads, new root production peaked amount of water required to return the soil
at bloom and again postharvest, with little to field capacity (drainage lysimeter).
growth in midsummer. Clearly, the seasonal Although there have been several lysimeter
pattern of root production is rather plastic. studies on apple, quoting actual totals for
These patterns of growth have not been corre- water use is not helpful, as the rates of water
lated with water status or nutrient uptake, use varied depending on the climate where
however. In general, Atkinson (1980) con- the work was done and the unique combina-
cluded that new fine roots are not required for tions of tree characters and environment,
adequate water and nutrient uptake by apple which are typically not reported. These fac-
trees. There needs to be much more research tors will be reviewed below.
integrating root and top growth with water A useful expression is the water-use rate
relations and nutrient uptake. per unit leaf area in midsummer. In the mild
sub-humid climates of The Netherlands
(Kodde and Kipp, 1990), Germany (Chen and
8.4.4 Mycorrhizae and water uptake Lenz, 1997; and calculated from data in
Braun et al., 2000), New Zealand (Green and
Apple roots have vesicular arbuscular mycr- Clothier, 1988; Green et al., 1995; Mpelasoka et
orrhizal (VAM) associations, which probably al., 2000b) and New York State (A. Lakso,
play an important role in extending the sur- 1997 and 2001, unpublished data), it has been
face area for absorption of immobile nutri- found that water-use rates on sunny days are
ents, such as phosphorus (Trappe et al., 1973; approximately 1–1.7 l m−2 of leaf area. In the
Plenchette et al., 1982; Gnekow and south of France, Angelocci and Valancogne
Marschner, 1989). Good growth of apples (1993) found in July that mid-season rates
appears to depend on the VAM association, were correlated to leaf area but could exceed
but apples appear normally to have VAM in 2.5 l m−2 leaf area day−1 in arid climates.
the field. There has been a report that the
water uptake and water status of apple
seedlings in sterile medium were improved 8.6 Factors that Affect Water Use in
by inoculation with VAM (Runjin, 1989). Apple Orchards
How important the mycorrhizae are to water
uptake and status of mature apple trees in the Water use by apple orchards varies with tree
field is not clear. Koide (1993) concludes that characteristics and climate. The principles reg-
non-nutritional mycorrhizal effects on water ulating water use by crops have been studied
relations are minimal; however, Auge (2001) over many years and they provide the basis
concludes that there may be direct effects in for understanding the variations observed
many plants as well as nutritional and plant (Allen et al., 1998). In short, ET by crops is ulti-
size-related effects. Considering the very low mately driven by energy from solar radiation,
root-length density of apples and the likeli- but it is modified by temperature, vapour-
hood of localized drying in the root–soil pressure gradient (VPG), boundary layers of
interface, it seems likely that mycorrhizae still air and conductance of the crop.
could improve water relations by improving
the root–soil contact. This is another area that
needs further research to clarify. If we under- 8.6.1 Energy, radiation and humidity
stand more we may be able to vary soil and
management practices as they have been Transpiration of plants is driven by energy
shown to affect mycorrhizae in crops (Barea from solar radiation, which heats the air and
et al., 1993; Dorgo et al., 1997). exposed surfaces, such as soil, water and
174 A.N. Lakso
leaves. Although we generally consider VPG 20
Canopy transpiration (µmol s–1)

of the air, or from leaf to air, as a driving
variable, transpiration is also based on the 15
energy from solar radiation. This energy pro-
vides the potential for ET and may be
received directly by the orchard or may be 10
imported into the orchard by the movement
of outside air of varying temperature and 5
VPG. Although the energy determines the
potential for water use by trees, several other 0
important factors affect the actual water use. 0 10 20 30 40 50 60
Leaf area removal (%)
Fig. 8.3. Effects of reducing leaf area with varying
8.6.2 Leaf area severity of summer pruning on the canopy
transpiration rate of apple trees as measured with
The amount of leaf area on a tree is important whole-canopy chambers under sunny conditions
to its water use since the leaves provide the (K.-T. Li and A. Lakso, 2001, unpublished data).
most active transpiring surfaces and they also
intercept the radiation that drives transpira-
tion. As the leaves intercept radiation, the Beyond the role of leaves in intercepting
energy warms the leaves and provides the radiation, interior shaded leaves of trees still
energy for the evaporation of water within transpire, although at lower rates than
the stomatal cavities of the leaves. exposed leaves. This suggests that trees with
Consequently, water-use rates vary over the many shaded leaves will have lower WUE,
season with the development and loss of the since Marangoni et al. (1992) found that the
leaf canopy and the related radiation intercep- shaded leaves with less than 10% of full light
tion. A similar effect occurs over the develop- had only 10% of maximum photosynthesis
ment of the life of the orchard as the canopies but 50% of maximum transpiration. The
fill their space and intercept more radiation. A transpiration rate may not be valid, due to
good correlation of apple tree water use to the artificial conditions of the leaf chamber,
leaf area has been found (Wibbe and Lenz, but this author has found that the leaf con-
1989; Angelocci and Valancogne, 1993), ductances of shaded leaves may be up to
although there was probably also a correla- 60% of that for exposed leaves. It has been
tion of leaf area to radiation interception. shown in grapevines with whole-canopy
A recent study in our laboratory has chambers that removing about 25% of the
shown that reductions in leaf area and radia- total leaf area, but only from the inside of the
tion interception due to summer pruning of canopy, had little effect on canopy photosyn-
mature apple trees also reduces water-use thesis, but reduced transpiration by about
rates (Fig. 8.3). The reduction in transpiration 10% (S. Poni, personal communication).
rates was less than the reduction in leaf area,
since the pruning increased the proportion of
leaf area that was exposed and reduced the 8.6.3 Crop structure and boundary layers
light interception less than the leaf area. The
lower transpiration of the pruned trees conse- The tall structure of apple trees, especially in
quently translated into less negative stem rows, makes a very aerodynamically rough
potentials. In drought years such summer structure. Consequently, air mixes in the
pruning may provide a method to improve canopies very well, so that the leaves are well
the water status of unirrigated trees without coupled to the environment. Thick, still bound-
affecting soil moisture. Severe pruning of ary layers of air, which can trap transpired
peach and pear trees has been demonstrated humidity over low crops like grasses or cover
to reduce extreme drought effects (Proebsting crops, do not develop over the canopies of
and Middleton, 1980). apple trees (Jarvis, 1985). Because of the excel-
lent mixing of bulk air with the crop, the leaves LSD
are exposed to the bulk-air humidity. This 5%
600
means that radiation and bulk-air VPG are
Total water consumption per plant (l)

both important environmental regulators of
water use in apples, along with stomatal con- 500
ductance. This will be discussed further later.
400
8.6.4 Factors affecting stomatal opening
300
Since stomatal opening has an important role
in regulating apple tree transpiration, factors
that affect stomatal conductance are of 200
importance. In apple trees in the field, it
appears that the stomata are well coupled to
photosynthesis, usually not opening more 100
than needed to maintain a constant internal
CO2 (Lakso, 1994). This means that factors
affecting photosynthesis will also affect tran- –Fr +Fr ++Fr
spiration. Crop load has been shown to posi- Fig. 8.4. Total water consumption of apple trees
tively affect gas exchange and water-use over 3 years if the trees never carried a crop (−Fr),
rates in several ways. Stomatal conductance carried a crop in the third year (+Fr) or carried crops
and photosynthesis of leaves are reduced as in the second and third years (++Fr). Leaf areas per
very low or zero crop loads are reached tree were 4.9, 3.4 and 2.5 m2, respectively.
(Palmer et al., 1997); thus non-cropping trees (Reproduced from Lenz, 1986, with permission.)
use less water per unit of leaf area (Hansen,
1971; Lenz, 1986; Navara, 1987; Wibbe and
Lenz, 1989; Buwalda and Lenz, 1992; Increasing planting density has also been
Masarovicova and Navara, 1994; Blanke, shown to increase water use (Atkinson, 1981).
1997; Chen and Lenz, 1997). If, however, the
cropping reduces leaf area more than it stim-
ulates the transpiration per unit leaf area, the 8.6.6 Interactions with drought
total tree water use may decline (Fig. 8.4).
As drought develops in apple trees, there are
several responses that affect orchard water
8.6.5 Canopy form, spacing and light use (Landsberg and Jones, 1981; Jones et al.,
interception 1985; Lakso, 1994). A reduction in vegetative
growth due to early-season stress will reduce
The canopy form and spacing of apple trees leaf area and possibly canopy light intercep-
can have a significant effect on water use by tion. Crop load may be reduced by early
orchards, with wider or larger tree forms stress (Powell, 1974), leading to lighter crop
using more water than thinner or more verti- loads and the related effects. These responses
cal forms. This effect is probably related to can cause adjustments in water requirements.
the varying effects of training systems on Of course, stomatal closure is important, as
light interception and secondarily on leaf well as leaf abscission in extreme cases.
area. Broader canopies, such as Y- or V-
shaped trees, or orchards that are spaced
more closely will intercept more light than 8.7 Water Deficits and Apple-tree
narrower, more widely spaced forms Growth, Cropping and Physiology
(Jackson, 1980; Jones et al., 1985; Palmer, 1989;
Robinson and Lakso, 1991; Lakso, 1994) and If the water-use demands of a tree cannot be
will use more water (Chen and Lenz, 1997). met, stress will develop. The effects of water
176 A.N. Lakso
stress on apple-tree growth and function water potentials indicate that shoot expansion
have been reviewed in depth previously by is almost linearly reduced by declining mid-
Landsberg and Jones (1981), with additional day stem water potentials (Fig. 8.5). Although
reviews more recently (Jones et al., 1985; mature leaves can osmotically adjust to main-
Lakso, 1994; Behboudian and Mills, 1997). tain turgor, apple shoot tips do not (Lakso et
This review will summarize the main conclu- al., 1984). Therefore, shoot-tip turgor and
sions from these reviews and integrate more growth will decline directly with declining
recent information. water potentials. Fruits and roots have been
shown to adjust osmotically for turgor main-
tenance (Beruter, 1989; Failla et al., 1992; Wang
8.7.1 Timing of stress et al., 1995; Mills et al., 1997). Again, the evap-
orative demands on the canopy will accentu-
A general observation is that processes that ate or ameliorate the effects of changes in soil
involve growth by expansion and especially water potentials by affecting the water poten-
those involved in growth by cell division are tials during the afternoon when it appears
more sensitive to water stress than processes that apple shoots grow (Powell, 1976a). Very
such as cell expansion, storage and gas similar responses have recently been docu-
exchange (Hsiao, 1973). Consequently, water mented for peaches, which also grow primar-
stress that develops in the spring and early ily in the late afternoon and evening (Berman
summer can have dramatic effects on vegeta- and DeJong, 1997a,b).
tive growth, fruit growth and fruit set, Root growth in a rhizotron has been
because early-season shoot growth and early shown to be reduced in response to drying
development of fruits are primarily by cell- soil (Rogers, 1939), although compensatory
division processes (Powell, 1974; Ferree and growth may occur in the wet-soil zones. Over
Schmid, 1990). Water stress that develops several months of rain-shielding, Jones et al.
more typically in midsummer will have less (1983) found that the shielded trees produced
effect on vegetative growth and less effect on more root length than either the rain-fed con-
fruit yield, as canopy development and fruit trols or shielded trees with added irrigation.
set are complete or nearly so by midsummer. Goode et al. (1978), however, found more sur-
Postharvest water stress has not been exam- face roots under irrigated trees but no effects
ined extensively, since there is the general
feeling that it is too late to significantly affect
1
physiological and growth processes. This
Control
may not be true in long-season climates, Stress
Shoot growth rate (cm day–1)
since there may be several months of good

weather and active physiological processes 0.75
before leaf fall. Late-season processes, such
as flower-bud development, root growth and
nutrient uptake, reserve storage and winter 0.5
acclimatization, would be expected to be
affected by stress, although water-stress
effects late in the season in warm climates 0.25
need more study (Kuroda et al., 1985).
0
8.7.2 Vegetative growth
–2.5 –2 –1.5 –1 –0.5 0
Since adequate water is needed for the turgor Stem water potential (MPa)
to drive expansive growth of apple leaves Fig. 8.5. Relationship of extension-shoot growth
(Davies and Lakso, 1979b), shoot growth is rate to variations in midday stem water in apple
sensitive to water deficits. Detailed measure- trees as affected by drought stress (M. Al-Hazmi and
ments of shoot growth rate in relation to plant A. Lakso, unpublished data).
deeper in the soil. Although individual roots Water stress reduces several aspects of
may slow growth in relation to drying soil, fruit growth and development. Fruit set in
the behaviour of the whole root system is the first weeks after bloom appears to
probably much more complex, as it responds depend on maintenance of an adequate rate
to variable soil moistures in the field. Several of fruit growth. Therefore, reductions in fruit
studies have shown that in arid climates over growth during the early cell-division period
time apple roots tend to concentrate under can reduce both fruit set and the potential for
drip emitters (Huguet, 1976; Levin et al., 1979, good fruit size at harvest (Powell, 1974),
1980; Crew and Funk, 1980; Bravdo et al., although often these early-season processes
1992; Neilsen et al., 2000). are completed before severe stresses
It should be noted that, due to the strong develop. Reductions in fruit growth are the
relationship of water potential to transpira- most common fruit responses to water stress
tion (Fig. 8.1), the severity of any soil-mois- in apples (Lord et al., 1963; Guelfat’Reich et
ture stress on leaf behaviour or growth will be al., 1974; Assaf et al., 1975, 1982; Goode et al.,
accentuated by higher evaporative demands 1978; Lötter et al., 1985; Ebel et al., 1993; Kilili
and, conversely, ameliorated by cooler, lower- et al., 1996b; Mills et al., 1996). The effects of
demand conditions. Consequently, as the soil water stress on fruit development appear to
moisture declines, stomatal opening and be more severe if the stress occurs during the
water loss will be maintained longer if the cell-division period compared with during
weather is cool and humid than if it is hot and the cell-expansion period. Reductions in
dry. A related factor is that slower stress growth during cell division are manifested
development allows apple leaves to osmoti- over the remainder of the season, even if
cally adjust markedly, which allows stomata water is abundant later (Fig. 8.6). The reduc-
to stay open longer than expected (Goode and tion in fruit size caused by water stress may
Higgs, 1973; Lakso, 1979; Lakso et al., 1984; cause firmness to increase. In some cases,
Jones et al., 1985). this increase in firmness has been found to
be independent of fruit size (Mpelasoka et
al., 2000a), while, in several cases, the
8.7.3 Fruit growth, quality and postharvest increase was not significant when compara-
effects ble fruit sizes were compared (Lord et al.,
1963; Ebel et al., 1993).
The effects of water stress on other fruit- There are many apparent contradictions
quality and postharvest characteristics are in the many reports of water stress on fruit-
complex and variable, since water relations quality effects, but several general trends
affects so many plant processes. In his occur. Besides the strong effect on fruit size,
review Sharples (1973) stated: increases in fruit dry matter or per cent sol-
It is seldom possible to separate the effects on uble solids have been quite consistent (Assaf
the fruit which are due to water supplies alone. et al., 1975; Ebel et al., 1993; Kilili et al., 1996a;
For example, variable levels of irrigation Mpelasoka et al., 2000a). Mills et al. (1994)
influence the overall nutrition and growth of found the effect of water stress on dry matter
the tree, and any observed effects on fruit and per cent soluble solids to decrease with
storage quality may be due as much to these later harvests, suggesting late-season sugar
general influences on the tree as to any specific development. Starch degradation appears to
influence of the water supply to the fruits.
be delayed by water stress (Powell, 1976b;
A common effect is the increase in vegetative Ebel et al., 1993), but generally water stress
growth with irrigation, leading to denser has led to earlier ethylene production (Lord
canopies and more shaded fruits, which et al., 1963; Ebel et al., 1993; Mills et al., 1994;
show effects of both water and shade. Also, Mpelasoka et al., 2000a). Water-core is more
early stress that leads to reductions in fruit commonly related to maturity so differences
set causes many crop load-induced differ- in water-core may be expected in relation to
ences in fruit quality, making differentiation the above effects on maturity (Marlow and
of causes difficult. Loescher, 1984).
178 A.N. Lakso
200
Control 1995
Stress 1995
Control 1994
150 Stress 1994
Estimated fruit weight (g)
100
Rewatering
50 95 94
0
0 20 40 60 80 100 120 140
Fig. 8.6. Apple fruit growth as affected by short-term water stress during cell division (1995) or after cell
division (1994) on seasonal fruit development (M. Al-Hazmi and A. Lakso, unpublished data).
The incidence of fruit disorders is quite 8.7.4 Gas exchange

variable in relation to water stress. Scald has
been reported to be both increased in dry There have been a great number of studies of
years but also decreased due to smaller fruit water stress on apple leaf photosynthesis
size caused by water stress (Wilkinson and and transpiration. The responses are typical
Fidler, 1973). Lötter et al. (1985) found that of any crop in that with loss of turgor, stom-
scald was worse with early-season stress but ata close and photosynthesis declines
decreased by late-season stress. Bitterpit and (Landsberg and Jones, 1981; Jones et al., 1985;
corking disorders have been reported to be Lakso, 1994). An important characteristic of
both increased and decreased (Sharples, 1973; apple water relations is that mature leaves
Goode, 1975; Lötter et al., 1985). Since most can osmotically adjust by as much as 2 MPa
calcium uptake into fruit occurs in the first or more over time as stress develops in field
several weeks of the growing season, it is trees (Goode and Higgs, 1973; Lakso et al.,
likely that different timings of water stress 1984). The osmotic adjustment in mature
may have different effects. Later-season stress leaves is primarily due to an accumulation of
that reduces final growth may reduce the monosaccharides, especially sorbitol (Wang
‘dilution’ of calcium concentration, so that the and Stutte, 1992; Wang et al., 1995). The
smaller fruits at harvest will have higher con- adjustment allows for turgor maintenance,
centrations even though the total calcium per which helps maintain gas exchange longer
fruit may be similar. This effect was noted in a into a drought. Chlorophyll fluorescence has
study of late-season European red-mite reduc- also been examined during water stress.
tions of fruit size leading to fewer calcium- Generally, it appears that initial or short-
related disorders (Francesconi et al., 1996). term stress responses are primarily related to
Clearly, a more fundamental understanding of gas exchange, but that, with longer drought
these interactions of calcium, fruit growth and periods, photochemical quenching is affected
crop load is needed to be able to interpret the (Jones et al., 1990; Massacci and Jones, 1990;
complex results from field studies. Fernandez et al., 1997b). Research has shown
that roots of young apple plants in drying there is a transpiration-induced adjustment

soil appear to signal the top of the plant to in water potentials (see Fig. 8.1). To reduce
reduce transpiration and leaf growth as if radiation absorption and leaf area, apple
there was water stress, even though ade- trees initially reduce extension shoot growth
quate water is available from the wet roots directly with increasing stress. With further
(Gowing et al., 1990). How important such stress development leaves may eventually
mechanisms are in the field still needs to be abscise, although mature leaves can osmoti-
determined. cally adjust significantly to maintain turgor
and function through moderate stress peri-
ods. To maintain water uptake with decreas-
8.8 Integration of Water-stress Effects ing soil moisture or higher evaporative
demand, apple trees have been shown to
Since most studies of water stress in apples increase root hydraulic conductivity
have focused on only a few aspects, it is use- markedly in response to greater evaporative
ful to attempt to integrate the overall effects, demands (Davies and Lakso, 1979a). Root
based on the format of Turner (1986). It eval- growth declines in dry soil, but roots
uates avoidance mechanisms, tolerance by develop further in the wet zones of the soil,
maintaining a high water potential, tolerance adjusting the water uptake pattern to opti-
to low water potentials and finally tolerance mize the use of soil water resources. Green et
to desiccation, which is rare in apple produc- al. (1997) have also found that individual
tion and therefore will not be considered. roots can adjust their water-uptake rates to
be able to maintain total water uptake from
those roots in wet soil.
8.8.1 Drought avoidance or escape
The perennial nature, low-temperature 8.8.3 Tolerance of low water potentials

threshold for spring growth and rapid
rosette-type leaf development allows for sig- The primary mechanism for tolerating low
nificant canopy development before most water potentials is the maintenenace of tur-
stress can develop. This is important, as gor. Turgor maintenance in response to
many early-season processes, such as initial declining water potential requires osmotic
fruit and shoot growth, are very sensitive to adjustment to counter the decreasing total
water stress. water potential. As water potential becomes
more negative with drought stress, leaves of
apple can lower the s (i.e. it becomes more
8.8.2 Tolerance by maintaining high water negative) by accumulating sugars and other
potentials solutes to maintain turgor and leaf function
(Goode and Higgs, 1973; Lakso et al., 1984;
The primary mechanisms of maintaining a Jones et al., 1985). Apple fruits also accumu-
high water potential within a plant are to: (i) late many solutes during development,
reduce water loss by stomatal regulation, which affect the fruit s and fruit water rela-
reduction of radiation absorption and reductions. Additionally, the hydrolysis of starch
tion of leaf area; and (ii) maintain water to sugars as the fruit matures lowers the s
uptake in spite of declining water availabil- (i.e. it becomes more negative) without
ity by increased root growth and increased requiring imported carbohydrates.
root hydraulic conductivity. Apple trees Consequently, the overall strategy for apple
maintain high water potentials in both ways. response to water stress appears to be to
The reduction in water loss in apple is reduce or stop leaf-area development (shoot
accomplished by very good stomatal regula- tips do not osmotically adjust), maintain good
tion, as discussed earlier. This leads to good WUE with stomatal coupling to photosynthe-
overall WUE. As stress develops further, sis and tolerate additional stress with osmotic
stomatal closure reduces transpiration and adjustment of the mature leaves, fruits and
180 A.N. Lakso
roots. Only with rather severe stress do apple results have not been consistent. For exam-
trees abscise leaves. This strategy would ple, M.9 has been reported to be less affected
appear to be best suited to shorter droughts or by drought than other stocks (Giulivo et al.,
very deep rooting where some water would 1985; Fernandez et al., 1997a; Kaynas et al.,
always be available. To make such a strategy 1997); however, other studies have come to
effective, the function of the leaves that remain the opposite conclusion (Chandel and
and osmotically adjust must persist to main- Chauhan, 1990). Finally, Alleyne et al. (1989)
tain canopy function, since new leaves are not found differences in diurnal stem potentials,
continually produced. Apple leaves do have but the differences were not related to stock-
an exceedingly long functional life. Several size category. Rogers (1939) summarized rhi-
studies have shown that the photosynthesis zotron observations and suggested that root
of healthy, exposed leaves is relatively stable growth slowed when soil water potentials
for at least 5–7 months if the environment fell below about 50 kPa; unfortunately, soil
allows (Porpiglia and Barden, 1980; Fujii and water potentials are rarely reported.
Kennedy, 1985; Wünsche et al., 2000). Such Physiological approaches have suggested
long leaf functional life allows the apple tree that a good drought tolerance was correlated
to maintain productivity with water stress with higher levels of proline, abscisic acid
while not producing new leaves continu- and carbohydrates in the leaves of the scion,
ously. In orchard conditions where extreme which were related to rootstock and soil
stresses are rare, this appears to be effective moisture (Chandel and Chauhan, 1991).
at maintaining productivity over a range of Again, the actual scion water status was not
stress severity. reported. Atkinson et al. (1999) found that all
rootstocks they tested produced about 42 m
of fine roots g−1 of root dry weight, but that
8.9 Effects of Rootstocks on Apple Water dwarfing capability was not correlated with
Relations and Drought Tolerance root production Additionally, several of the
stocks (including M.9, M.26 and MM.111)
The effect of rootstocks on water relations produced more fine roots in response to
would seem to be important and yet over drought stress, while M.27 and several AR
many years of study and observation a clear stocks generally produced fewer roots, but
picture has not evolved. This is probably due the responses to drought were not related to
to the wide range of direct and indirect fac- vigour control. Psarras and Merwin (2000)
tors that may be involved. Consequently, the found a slight shift towards finer root diame-
review of apple rootstocks by Ferree and ter with water stress of M.9 and MM.111, but
Carlson (1987) only stated that MM.111 is total root dry-matter production declined
reported to be the most tolerant to drought, strongly and root respiration increased with
while M.9 and M.26 are reported to be sensi- soil moisture stress.
tive. A recent guide for rootstocks made little
mention of drought resistance (Wertheim,
1998). Certainly, differences in tree response 8.9.2 Water-relations studies in the field
to water stress related to rootstocks can be
observed in field situations. The difficulty The physiology and growth responses to
lies in understanding the bases of these normal field conditions or to drought stress
responses and whether they represent direct in the field of trees on different rootstocks
physiological effects or indirect effects. have also been evaluated. Some have found
no or few differences in response to drought
or irrigation by rootstock (Ferree and
8.9.1 Controlled drought-response studies Schmid, 1990; Higgs and Jones, 1990).
Shorter-term studies of diurnal patterns of
There have been several studies utilizing stem potentials showed that non-stressed
potted trees of varying age to give controlled apple trees on M.9 and M.26 had more nega-
water-stress treatments. Unfortunately, the tive midday stem potentials than on M.7,
MM.106 or MM.104 (Olien and Lakso, 1986). was attained. Potted-tree studies may induce
Calculated hydraulic conductivities of xylem variable responses to water withholding that
water transport suggest that rootstocks differ are due primarily to variable depletion of a
in their ability to conduct water to the scion. constant pot soil volume by trees with differ-
Tree size was well correlated with midday ent leaf areas. Considering all the complexi-
stem potentials in this group of stocks, and ties mentioned above, it would be desirable
shoot expansion rate was directly related to for future studies to relate responses to esti-
midday stem potentials, which decreased mates of either soil or plant water status,
with increasing vascular distance from the such as predawn water potential or midday
roots (A. Lakso, 1984, unpublished results). stem potential (Olien and Lakso, 1986;
Alleyne et al., 1989). This will help to differ-
entiate the effects of stress due to root distri-
8.9.3 Indirect effects bution or plant size (in relation to soil
volume) versus internal effects, such as
Apple rootstock root systems have very low hydraulic conductance or hormone balances.
root-length densities (root length per volume This author proposes that, under field
of soil), have non-uniform distributions and conditions with a non-restrictive soil, there is
differ in their responses to soil structure probably a greater midday water stress
(Rogers, 1939; Fernandez et al., 1995). induced under high transpiration rates in the
Consequently, root distribution and density more dwarfing stocks, due to lower
patterns as affected by rootstock in any given hydraulic conductivity compared with the
soil will determine the total volume of more vigorous stocks. As the summer
potentially available soil water and deter- becomes warmer, the shoot growth of the
mine the availability of water independently more dwarfing stocks terminates earlier (R.S.
of any direct physiological response of the Johnson and A. Lakso, 1982, unpublished
rootstock itself. Rootstock interactions with results). For the more vigorous stocks, shoots
scion growth and cropping also complicate can then grow more until the length of the
such studies. Rootstocks differ in their ten- branches reaches a length where the total
dency to induce cropping, especially early hydraulic resistance (the rootstock plus the
cropping in young orchards (Wertheim, resistance of flow in the branch) is similar to
1998). As discussed earlier, cropping has that at the end of the shorter branches on the
strong effects on water use per unit leaf area, dwarfing stocks. This would explain why the
and the growth of the root system is strongly summer leaf water potentials and stomatal
inhibited by heavy crops. Hewett and conductances are generally similar for exte-
Cassidy (1977) and Goode et al. (1978) found rior leaves of a scion on many different root-
that irrigation gave stronger effects in heav- stocks (as also noted by Ferree and Schmid
ily cropping trees (e.g. the ‘on’ years of a (1990) over many rootstocks). It may also
biennial orchard). explain why apple trees, and other tree
Finally, many studies have been done species, tend to stabilize at a spherical form,
with young potted rootstocks or trees where all exterior shoot tips are similar dis-
grafted on different rootstocks. Since root- tances from the root crown.
stock breeders feel that at least 6 or 7 years of
growth in the field are required to establish a
stable ranking of rootstock vigour, a correla- 8.10 Management of Water Relations in
tion of mature field behaviour with that of the Field
young potted trees may not necessarily be
expected. Indeed, many of the studies 8.10.1 Orchard water balance
reviewed reported plant-size rankings that
were not in accordance with mature size To manage the water balance of an apple
ranking. Also a ‘drought treatment’ only orchard requires knowledge of water use
means that water was not applied; it does over the season in the given climate, the
not mean that the same plant water status water status needed for the desired tree per-
182 A.N. Lakso
formance, and other factors affecting the 8.10.2.2 Soil-moisture monitoring

water balance of an orchard (rainfall, soil
Another common approach is to monitor
reserves, climate, cover crops and cultivar).
soil moisture with various soil-moisture
In arid climates, the rainfall and soil reserves
devices and relate soil moisture to plant
are generally so deficient compared with ET
water use and plant stress (for reviews of
demands that the decision to irrigate is clear.
measurement of soil moisture, see
However, in humid climates, lower ET
Campbell, 1988; Ley, 1994a; Hanson and
demands and greater rainfall totals and vari-
Peters, 2000; Hanson et al., 2000). Since the
ability make it difficult to determine the eco-
soil water content is measured, these meth-
nomic feasibility of irrigation. In this case the
best approach is a risk assessment for water ods are good for determining soil water
limitations. This should include the soil depletion, which is needed to estimate
reserves (soil rooting volume, soil water- whole-orchard ET and irrigation needs.
holding capacity and initial water content in The greatest general limitation with soil-
the spring), the average rainfall, the average moisture monitoring is sampling in hetero-
tree demands and the demands of any com- geneous soils in which there are extremely
peting plants such as cover crops between low root densities and erratic apple root dis-
the rows (common to reduce erosion in high- tributions. Even in quite uniform soils, it is
rainfall areas). In some cases adequate water difficult to determine where to place a lim-
status may be obtained in dry seasons sim- ited number of sampling sensors or access
ply by reducing water use of competing sites, since it is difficult to know where the
plants with herbicides. A quantitative risk roots are located, especially for dwarfing
analysis is very useful; however, it is beyond rootstocks. Due to this problem, soil-
the scope of this review. moisture monitoring is best suited to
developing orchard water-use estimates with
soil water-balance studies, rather than to
8.10.2 Irrigation management estimating plant water status.
Irrigation is primarily to provide supplemen- 8.10.2.3 Evapotranspiration estimations

tal water not provided by rainfall or soil
water reserves. Consequently, efficient irriga- Another common approach is to estimate
tion management requires knowledge of the the environmental demands for ET as modi-
water loss of the apple orchard (trees, soil fied by orchard architecture and plant resis-
evaporation and cover crops/weeds) and the tances for water vapour. There have been
soil water reserves and rainfall. There are many studies of microclimatology and the
several practical approaches used to estimate factors that drive ET and many equations
water usage or tree water status for schedul- developed to estimate ET. The recent Food
ing irrigation applications. and Agriculture Organization (FAO) book
(Allen et al., 1998) is an extensive review of
this area and is recommended for study,
8.10.2.1 Experience
while Hanson et al. (1999) is more grower-
A very common qualitative approach is a orientated.
‘mental model’, which is essentially a risk In general, the major environmental fac-
assessment based on grower experience with tors are net radiation (Rn), humidity and
the lengths of droughts and evaporative temperature, which lead to vapour-pressure
demands in relation to the perceived soil deficits (VPD), and wind speed, which
water reserves and their irrigation tools. affects the boundary-layer resistance
Although not quantitative or documented, a around the crop. Crop characteristics gener-
good grower is able to take into account a ally affect ET by affecting the boundary-
wide range of unique characteristics of their layer resistance (canopy height,
individual orchards that general models can- aerodynamic roughness and density), the
not accommodate. radiation energy balance (reflectivity and
ground cover or light interception) and the 1.2
Transpiration, E1 (mm h–1)

stomatal resistance to water loss. There are
important interactions among these factors.
0.9 Forest
These factors have been summarized in sev-
eral models of ET, which are similar in that
they incorporate these main factors with 0.6
various simplifying assumptions. The Field crops
model most commonly used is the original
or modified Penman–Monteith equation 0.3 Short grass
(Allen et al., 1998).
One important characteristic unique to 0
tall, discontinuous canopies like apple 0 10 20 30 40 50
orchards is that the tall, rough canopies Canopy conductance, g L (mm s–1)
cause air turbulence, which reduces crop
boundary-layer resistances. This means that Fig. 8.7. The general response of crop transpiration
the tree canopies are well coupled to the bulk to canopy conductance comparing low grass, field
atmosphere above (Jarvis, 1985). crops and forest (from Jones, 1992, with permission).
Consequently, in apple trees the stomatal
resistances are important regulators of ET, has been estimated from fewer studies, but
since transpired water vapour quickly moves apple ET has been related to the grass refer-
to the bulk air and the VPG is not affected. ence ET by a scaling factor called a crop
This is different from low, smooth, continu- coefficient (Kc) (Ley, 1994b; Allen et al.,
ous crops like grasses, which can develop a 1998; Hanson et al., 1999). Weather data are
thick boundary layer that traps humidity. If used to calculate the reference grass ET,
the grass stomata open, the transpired water which is then multiplied by a crop coeffi-
vapour cannot escape, so the VPG declines, cient to adjust for crop differences:
counterbalancing the opening of the stomata.
Thus, grass-crop ET may not respond signifi- Apple ET = ETgrass × Kc apple
cantly to changes in stomatal resistances Similarly versus Epan:
(McNaughton and Jarvis, 1983; Jarvis, 1985;
Apple ET = Epan × Kc apple/pan
Jones, 1992; Fig. 8.7). Consequently, apple
tree ET is controlled by Rn, VPG and stom- Basal crop coefficients have been esti-
atal conductance, while grasses or cover mated with the assumption for fruit-trees of
crops between the rows of apples respond 60% ground cover, no water stress, a large
primarily to Rn (Plate 8.1). continuous orchard and a moderate climate
(Allen et al., 1998). There are several correc-
tions or adjustments that have been devel-
8.10.2.4 Crop coefficients for irrigation
oped that are appropriate for apple orchards,
Since complete ET analyses to determine though not easily applied due to the erratic
absolute ET cannot easily be done with all canopies and alleys (Allen et al., 1998).
crops in all climates, a method has been Corrections are made for the seasonal
developed where individual crop ET is development of the apple canopy and thus
related to a reference crop or to another variations in seasonal water use (Fig. 8.8).
common measure, such as class A pan Additionally, the crop coefficient for apple
evaporation (Epan). A healthy, low grass is orchards is affected by other factors that
typically the reference crop, since the ET of affect tree water use. First, due to the dis-
grass has been studied extensively and continuous canopies, ground coverage is
over extended periods and is well esti- used as a simplification for light intercep-
mated from meteorological data by the tion, the driving energy. Mature orchards
modified Penman–Monteith equation are assumed to be about 60% ground cover,
(Allen et al., 1998). This reference is called giving full ET (Fereres and Goldhamer,
ETo or ETgrass. In contrast, apple-crop ET 1990; Hanson et al., 1999). Apple orchards
184 A.N. Lakso
Evapotranspiration (mm day–1)

18
15
12
0
0 30 60 90 120 150 180 210 240
Days after bloom
Fig. 8.8. Seasonal pattern of apple-orchard water use (from Beukes and Weber, 1982, reproduced by
permission of the Journal of Horticultural Science).
vary in mid-season ground coverage, drought or other stresses (crop-load effects,

since so many canopy forms are used. for example); and (iv) differences between
This author has found light interception orchards with a cover crop between the rows
of mature commercial orchards to vary versus those with bare soil (Allen et al.,
from about 25% to about 80%. The mid- 1998). Clearly, it is not possible to provide
day ground-cover estimate has been simple guidelines to estimate the ET of apple
found to be useful for horizontal to fairly orchards, due to the myriad of climates, trees
natural round trees, but it underestimated and adjacent environments.
daily light interception for slender spindle Consequently, it is highly desirable to do
and thin, vertical palmette forms by 14 local studies of water use in each climatic
and 35%, respectively (Wünsche et al., region to determine local Kc values, rather
1995). than accepting Kc values from other regions.
Corrections are also based on differences
in humidity and wind in different climates,
and the differences are accentuated with tall 8.10.3 Plant-based methods for irrigation
crops, such as apple orchards (Allen et al., scheduling
1998). For example, in arid climates with low
humidity, the Kc for tall crops and for higher The many indirect methods discussed are
wind speeds must be increased as much as limited to general guidelines or values, since
30% for 5 m tall apple trees with 6 m s−1 there are so many unique combinations of
average winds. In contrast, in humid cli- environmental and soil conditions. These
mates with calm winds, the Kc must be may be adequate for scheduling irrigation,
decreased by 20–25%. This indicates, for but, in intensively managed orchards, more
example, that, for apple trees and other tall direct methods to allow the tree to provide
crops, Kc values developed in arid climates information on water needs have been
(where such studies are usually done) may desired. The bases of most of these methods
not be correct in more humid climates, have been reviewed earlier in the section on
where many apples are produced measurement of tree water status.
(Annandale and Stockle, 1994).
Additional adjustments that are made
8.10.3.1 Visual inspection
relate to: (i) edge effects, called ‘clothes-line’
or ‘oasis’ effects, for small plantings that are The most common method is visual inspec-
adjacent to different sizes or types of vegetation of the trees for signs of wilting, growth
tion; (ii) lower stomatal conductances than inhibition, leaf colour, etc. Although these
are normally assumed for the grass; (iii) symptoms indicate stress, usually they
reductions in stomatal conductance due to appear too late to make an early intervention.
8.10.3.2 Midday stem potential use demands of the trees (i.e. at a deficit).
The term regulated deficit irrigation (RDI)
The midday stem potential is a very good
implies the same concept, except perhaps
integrator of water status at any time and
with the targeting of specific growth stages
has been well related to important crop
versus full-season deficits; the general term
processes, as reviewed earlier. Although the
deficit irrigation will be used here.
midday stem-potential method is feasible for
Behboudian and Mills (1997) have recently
large growers with qualified technical sup-
reviewed deficit irrigation, so this discus-
port or consultants, the use of the pressure
sion will address how it may apply in
bomb is still considered too technical for
apple production.
most growers.
In addition to saving water, the use of
deficit irrigation at critical times during
8.10.3.3 Temperature monitoring crop development has been tested to con-
trol vegetative growth without harming
As discussed earlier, monitoring leaf tem-
fruit development (Chalmers et al., 1981) or
perature has been used to estimate when
to improve apple fruit quality (Mpelasoka
plants need water. However, the responses
et al., 2000a). A key point was to identify a
of apple stomata to other factors, such as
stage of development when the fruit were
crop load, may give false signals, (Jones,
not actively growing or not sensitive to
1994, 1999a,b). Also the dynamics of vari-
stress, but the shoots were still active. This
able radiation, VPD and wind makes it dif-
occurs in stone fruit and grapes, which
ficult to use temperature monitoring in
have a double-sigmoid fruit-growth pattern
many humid or cloudy climates. Jones
with a mid-season lag in fruit growth while
(1999a) has suggested using wet and non-
shoots are still vigorous. This scenario does
transpiring leaves as references instead of
not occur with apple, since the inherent
air temperature, but the dynamic variability
fruit-growth pattern by weight is best
and the cost of the instrument will limit its
described as ‘expolinear’ (exponential
use. Though not quantitative, the simple
early, then close to linear until normal har-
method of feeling the warmth of the largest
leaves in the sun with one’s fingers should vest) (Lakso et al., 1995). Although apple
be used more as an indicator of stomatal extension shoots may be sensitive to water
closure, as it is more sensitive than one may stress, fruit growth is also quite sensitive,
think and many trees can be checked easily especially in the early season when shoot
and quickly. growth is strong (Fig. 8.9). Therefore, man-
aging deficit irrigation to avoid loss of fruit
growth seems much more difficult in the
8.10.3.4 Trunk and fruit monitoring field with apples than with stone fruit or
As discussed earlier, the physical contraction grapes.
of the stems and fruit of apple trees has been Studies of deficit irrigation in apples have
examined as an integrator of water stress. shown that there are variable reductions in
This method is promising, but it needs cali- fruit weights with deficit irrigation in the
bration to determine thresholds for initiating field (Beukes and Weber, 1982; Lötter et al.,
irrigation and needs more rugged and inex- 1985; Ebel et al., 1993, 1995; Mills et al., 1994,
pensive equipment. 1997; Kilili et al., 1996a; Naor et al., 1997;
Behboudian et al., 1998; Mpelasoka et al.,
2000a; Fig. 8.10). Reductions in fruit size may
8.10.4 Deficit irrigation with apples not necessarily be detrimental if fruit size
normally tends to be too large for optimal
In many regions water may be limiting or quality. Conversely, reductions in fruit size
expensive, so there has been much interest may be doubly detrimental for heavily crop-
in improving efficiency of irrigation by ping trees, since fruit size is already reduced
using irrigation at levels below the water by the crop load.
186 A.N. Lakso
1.5 modelled, but no overall model has been

Control developed. Landsberg and colleagues began
Stress to include water relations in a general apple
Fruit growth rate (mm day–1)
model, but the model did not appear to have

1
been completed (Landsberg, 1980). Models
of stomatal response to the environment
have been developed (Landsberg and Butler,
1980; Thorpe et al., 1980; Jones and Higgs,
1989; Jones, 1998), with an emphasis on the
0.5 effects of VPD reducing conductance. This
has recently been expanded to scale up for
whole-tree transpiration estimates (Green
and McNaughton, 1996). Hydraulic flow and
0 capacitance of the tree have been modelled
–2.5 –2 –1.5 –1 –0.5 0 with electrical-circuit analogues (Landsberg
Stem water potential (MPa) et al., 1976; Jones, 1983). Perhaps the most
Fig. 8.9. Apple fruit growth rates as affected by useful modelling has been in regard to evapo-
varying stem water potentials induced by water transpiration of orchards (Butler, 1976;
stress (M. Al-Hazmi and A. Lakso, 1996, Thorpe, 1978; Thorpe et al., 1978; van der
unpublished data). Maas, 1992; Green and McNaughton, 1996),
which has used the energy-balance approach
(see Section 8.10.2.3). However, validation of
In conclusion, it appears that deficit irri- these models with independent data from
gation can be somewhat useful, but the different climates is needed.
apple growth habit is not particularly well Models allow quantitative hypotheses to
suited to this approach, especially in climates be developed that can be tested, but they
with summer rainfall. For control of vegeta- have limitations that must be considered in
tive growth, the early-season use of gib- relation to the unique characteristics of
berellin inhibitors, such as paclobutrazol or apple trees. Jones and Tardieu (1998) make
prohexadione-Ca, are probably better tools several important points relevant to model-
for that purpose. ling water relations of apples: (i) models are
best in homogeneous systems (apple
canopies and root systems certainly are not);
8.11 Needs for Integrative Approach to (ii) modelling water status in soils is difficult
Water Relations due to increasing spatial variability as the
soil dries; the low root density and clumping
From the preceding discussion, it is clear that of apple roots are a problem; (iii) modelling
a broad integrative view is necessary to be stomatal control of water relations is compli-
able to interpret results, especially from the cated by non-hydraulic factors that affect
field. The lack of adequate information on the stomatal conductance, such as crop effects in
environment, tree age, stage of tree develop- apple; (iv) rainfall interception by canopies
ment, crop load, soil type and related infor- may be an important component of orchard
mation was the major limitation in evaluating water balance and yet we lack information
the many publications in this area. on rainfall interception; and (v) uniform
canopies are easiest to model; apple canopies
have great variability due to various training
8.11.1. Modelling water relations systems. So modelling apple-tree water rela-
tions will be a difficult challenge.
Modelling is an approach that attempts to Finally, a precautionary note is needed.
quantitatively integrate the main controlling Most physiological studies of apple are done
factors in a system over time. There have to address applied problems of stress in the
been several aspects of apple water relations field and yet too often small, potted apple
Trickle
250
200
150
Hand-thinning
completed
100 Standard
Volume per fruit (cm3)
1.5 Fruit/TCA
50
4.8 Fruit/TCA
7.9 Fruit/TCA
RDI/Trickle
250 RDI
ended
200
150
Hand-thinning
completed
100
Standard
1.5 Fruit/TCA
50
5.4 Fruit/TCA
10.0 Fruit/TCA
17 June 15 July 12 Aug. 9 Sept.

Date
Fig. 8.10. Apple fruit growth over time of varying crop loads with full irrigation (top) and with mid-season
regulated deficit irrigation (from Ebel et al., 1995). The ‘standard’ curve is the expected curve for normal
crop levels in Washington State. TCA, trunk cross-sectional area in cm2.
trees are used as model systems under con- structural, physiological and cultural factors
trolled or semi-controlled conditions. Are they interacting. It is impossible to understand
a good model of the mature tree behaviour? these relations with a reductionist view,
In the case of water relations, there are so although detailed understanding of compo-
many differences in the environment of the nent processes is always valuable. We must
pot versus field soil and the tree behaviour strive to develop a broad integrative
that we cannot assume that potted trees are a approach in which many observations are
good model. We must accept the challenge of taken in concert with measurements.
working in the field with mature plants. Although more difficult experimentally, it
appears that we shall need to utilize plant-
based measures to integrate all the factors
8.12 Conclusions involved, since indirect measures, such as
soil moisture or climate, are inadequate.
In conclusion, the water relations of apple Midday stem water potentials are probably
trees are extremely complex and dynamic, the best single physiological measure of
with many soil, atmospheric, temporal, plant water status; however, better integrals
188 A.N. Lakso
of water stress are needed. Finally, the role water relations of apple trees will need to be
of modern molecular biology is potentially determined (Tyerman et al., 1999). It will cer-
exciting but not clear at this time. For exam- tainly require good teamwork and coopera-
ple, recent discoveries of aquaporins that tion across disciplines to evaluate how best
affect membrane hydraulic permeability to use these new tools to improve our
may be important, but the relevance to field understanding.
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9 Light Relations
Luca Corelli Grappadelli

Dipartimento di Colture Arboree, University of Bologna, Italy

9.2 Light: Physical Properties 196
9.3 Photosynthesis 197
9.3.1 Single leaves 198
9.3.2 Whole canopy 199
9.4 Source–Sink Relationships 201
9.5 The Basis of Orchard Productivity 202
9.5.1 Models 202
9.5.2 Orchard configuration studies 204
9.5.3 Light distribution 206
9.6 Light Effects on Physiological Parameters 208
9.6.1 Responses to light quality 208
9.6.2 Responses to light intensity 209
9.7 Controlling Light Levels in the Orchard 210
9.7.1 Bagging 210
9.7.2 Reflectants 211
9.7.3 Shading to reduce sunburn 212
9.8 Conclusions 213
9.1 Introduction tioning of resources (such as nutrients, car-

bon and water) into fruit instead of other
Visible light in the 400–700 nm waveband is organs are also largely controlled by light.
the driving factor of biomass production via Light therefore plays a twofold role in influ-
its effect on photosynthesis, so it is not sur- encing the processes that lead to the pro-
prising that production of dry matter in duction of large quantities of high-quality
apple has been shown to be related to the fruit: on the one hand, it supplies the
amount of visible light intercepted by trees energy stored in chemical form in carbohy-
(Palmer, 1989; Lakso, 1994). However, in drates; on the other, it influences the
fruit trees, including apple, dry-matter pro- ontogeny of the tree’s structures, so that the
duction does not automatically translate tree’s physiological traits are generally
into increased yield of marketable fruit. The enhanced in parts of the canopy where high
complex phenomena that result in the parti- light conditions prevail.
196 L. Corelli Grappadelli
This influence extends to many morpho- Visible light represents a fraction (about
logical and physiological traits of the vegeta- 42–45%) of the short-wave radiation spec-
tive components of a canopy, from bud trum released by the sun. Because photo-
differentiation to leaf attributes, to the ratio of synthesis is by far the most important plant
spur vs. shoot leaf area, to the photosynthetic phenomenon influenced by light, this por-
potential of these leaves and to the photosyn- tion of the spectrum is also termed photo-
thetic capacity of the different leaf types that synthetically active radiation (PAR). The
support fruit growth. Furthermore, the dura- definition photosynthetic photon flux (PPF)
tion of the light period and light intensity each is also appropriate, to indicate that the pho-
affect partitioning of carbon between different tosynthetic process responds to the rate of
chemical forms (transport or storage) and absorption of photons intercepted in this
between reproductive and vegetative sinks range and not to the rate of absorption of
and affect fruit quality attributes, such as skin energy. PPF has units of mol m2 s1
colour, soluble-solids concentration, acidity (micromoles of quanta of energy, i.e. of pho-
and the incidence of storage-related disorders. tons, per unit area and time). A typical PPF
Because of these many effects of light, value for the direct component of sunlight
knowledge of the basic relations between at noon on a clear day with the sun over-
light and the tree is necessary to manage head is approximately 2000 mol m2 s1.
the canopy in order to maximize its pro- In addition to direct irradiation from the
duction efficiency. This chapter is con- sun, two other light fractions contribute to
cerned with examining the main aspects of the total amount of energy available on the
tree light relations, beginning with a defini- ground: a fraction scattered by the clouds
tion of the main physical characteristics of and a fraction scattered by the atmosphere,
light, leading to an examination of several which together make up the diffuse irradia-
physiological aspects and then deriving tion component. On a cloudy day, diffuse
practical aspects of tree and crop manage- irradiation can be quite high relative to the
ment that have their foundation in this direct component. Because diffuse irradia-
scientific knowledge. tion can penetrate the canopy from virtually
every direction, the total light available in
inner tree-canopy positions will be affected
9.2 Light: Physical Properties by the relative amounts of the direct and
diffuse components: under western New
The irradiance (total radiant flux density) pro- York State conditions, Lakso and
vided by the sun that reaches the outer atmos- Musselman (1976) found that the quantity
phere, measured perpendicularly to it, is of light available at locations inside a
termed the ‘solar constant’ and its value is on canopy was increased under partly cloudy
average 1360 W m2. However, because of conditions, compared with that under clear
absorption by the atmosphere, the total radiant sky conditions (Fig. 9.1).
flux reaching the ground at sea level is about Latitude affects the average annual
58% of that value. If the whole spectrum of amount of light available daily – higher lati-
short-wave (wavelengths < 4 m) radiation is tudes have lower mean daily light integral
considered, the total energy available (global values. The disadvantage due to latitude is
irradiation) is approximately 800 W m2 at sea partly offset by the longer daytime duration
level. Global irradiation increases with alti- during the summer at the higher latitudes.
tude, although its actual values will depend on Both the intensity and the spectral distri-
clouds and atmospheric composition, since bution of light after it reaches the canopy
water vapour, CO2, gases and dust particles change dramatically within a short distance.
absorb and scatter radiation at different rates. Absorption by leaves averages about 80% of
Under clear sky conditions at noon, more than incoming visible light. A major portion of
1000 W m2 can be found at 2000 m above sea the ‘missing’ light is in the region of 550
level (Nobel, 1983), with a greater fraction in nm, where chlorophyll absorption is low,
the UV component than occurs at sea level. and a large fraction of this light is reflected
Light Relations 197
cloth, which, unlike natural within-canopy

Incident PAR (µmol m–2 s–1)
2000 shading, does not alter the light spectrum.

Clear
9.3 Photosynthesis
1000 Partly cloudy The major quantitative effect of light on

plants is on photosynthetic activity: in
properly managed orchards, increases in
photosynthesis (i.e. light intercepted) will
Overcast result in increases in yields of marketable
0 apples. The most light-efficient orchard con-
200 100 0 100 200 figurations have been reported as being
East West capable of intercepting 60–70% of available
Distance from tree centre (cm) radiation, which may translate into very
Fig. 9.1. The penetration of light within the canopy high yields – maximum yields reported are
of a 9-year-old ‘Golden Delicious’/M.2 tree, during from New Zealand, at 120–140 t ha1
3 days with different sky conditions. The greater year1, sustained over several seasons
penetration to the interior parts of the canopy (Lakso et al., 1999). Several authors have
occurred on the day with the highest diffuse radiation demonstrated the relationship between
component (the partly cloudy day). PAR, light interception and yield, as summarized
photosynthetically active radiation. (From Lakso and
by Lakso (1994), who also showed how, at
Musselman, 1976, reprinted with permission.)
light interception levels greater than 50% of
available light, orchard productivity may
or transmitted, leading to the green colour differ widely, from very poor to very high
of leaves. A rather large fraction of the yields. A partial reason why this may be so
energy load on the leaf is from infrared (IR) lies in the curvilinear relationship between
radiation from the sun or from the atmos- PPF and whole-tree photosynthesis (Fig.
phere, but it is in large part re-emitted at 9.2): in the absence of other limiting factors,
longer wavelengths (about 70% of the total when the light intensity available to the
irradiation is lost this way (Nobel, 1983)). canopy exceeds saturation (the light inten-
The spectral composition of the light within sity above which no further increases in
a canopy is thus changed, with a shift photosynthesis occur), the tree’s instanta-
towards longer wavelengths brought about neous potential is reached and no further
– in the visible range – by the selective photosynthetic gain will be obtained by fur-
absorption of the photosynthetic pigments ther increases in light interception.
(particularly effective in reducing the blue The light-response curve of a canopy, as
and red components) and by lower absorp- shown in Fig. 9.2, generally resembles that of
tion and greater reflectance in the IR. As a a single leaf. Depending on the canopy char-
result of this, the red/far-red ratio in the acteristics (size, density, degree of shading of
inner regions of the canopy is reduced, interior leaves by the external ones), however,
which may result in changes in phy- the values of the compensation point, the
tochrome-mediated responses. However, quantum efficiency and the saturation point
very little is known regarding the relevance (Flore and Lakso, 1989) for a single leaf and
of phytochrome in apple morphogenetic for a whole canopy may vary widely. The
responses. Indeed, many examples exist in greater the density of the canopy and the
the literature where responses generally amount of shaded leaves, the greater the dis-
attributed to phytochrome (leaf morphol- crepancy between single-leaf and whole-
ogy and photosynthetic characteristics, canopy measurements: in general, since the
flower-bud differentiation, fruit growth and whole-canopy gas exchanges normally
colour) have been induced by simply reduc- include fruit and wood respiration in addition
ing light levels with neutral-density shade to respiratory losses from the leaves, the com-
25
NCER (µmol CO2 m–2 s–1)

20
15
10
0
0 500 1000 1500 2000
–5
PPF (µmol m–2 s–1)
Fig. 9.2. Light-response curve of a 7-year-old ‘Golden Delicious’/M.27 tree, enclosed in a whole-tree
assimilation chamber. The tree was progressively shaded by adding layers of neutral-density shading cloth.
NCER, net carbon exchange rate; PPF, photosynthetic photon flux.
pensation point is higher for the whole versible: once a shade leaf is placed under
canopy. At the whole-canopy level, the inter- high photosynthetic photon fluxes (as a
action with light is more complex, as it inte- result of summer pruning, for example), it
grates many factors in the response, is not capable of reaching the same high
including, in addition to those outlined rates of fixation that can be reached by sun
above, tree shape, leaf density, crop load, leaves.
nutritional factors and water status. However, Sun leaves exhibit greater thickness
a whole-tree approach solves the problem of (related to the number of layers of cells in the
scaling-up individual leaf measurements, in palisade parenchyma (Doud and Ferree,
order to estimate a tree’s response. Because of 1980b)). They also have higher nitrogen con-
these differences, both single-leaf and whole- tent and are more dense (higher leaf-area
canopy responses are discussed. density (mg cm2)) than shade leaves (Flore
and Lakso, 1989). The photon fluxes during
leaf emergence and development at the
9.3.1 Single leaves beginning of the season influence these para-
meters, which, once set, do not vary consid-
Apple leaves exhibit a typical asymptotic erably. This constitutes one of the main
response of photosynthesis to PPF and their reasons for maintaining open canopies with
photosynthetic parameters – compensation, good light distribution throughout the sea-
saturation and quantum efficiency – are com- son. In addition to the effect of the current
parable to other major temperate fruit crops. light levels on leaf characteristics, some evi-
Apple-leaf morphology exhibits distinct dence exists that the light levels experienced
traits that identify the light environment by the bud during the differentiation process
under which the leaf differentiation in the previous season may influence para-
processes have taken place and which relate meters such as specific leaf area, at least for
directly to leaf productive potential. In fact, the primary spur leaves. Tustin et al. (1992)
leaves that have developed in the exterior, reported lower specific leaf area 2 weeks
well-illuminated parts of the canopy (‘sun’ after full bloom (AFB) in primary spur leaves
leaves) are capable of greater maximum pho- derived from buds that had differentiated
tosynthesis than leaves that have been under under low PPF in the previous season, even
low photon fluxes during their development though at 2 weeks AFB they were fully illu-
(‘shade’ leaves). However, shade leaves are minated (Table 9.1). In the same study, the
in general more efficient at utilizing low pho- bourse shoot leaves responded only to cur-
tosynthetic photon fluxes or the sun flecks rent light levels: severe shading (70% reduc-
that may occur in the internal portions of a tion) of leaves in high light positions
canopy. This differentiation is largely irre- significantly decreased leaf-area density
Light Relations 199
Table 9.1. Leaf area density (mg dry matter cm2) of spur and bourse shoot leaves growing under
different light regimes at 2, 5 and 8 weeks after full bloom, on 9-year-old ‘Golden Delicious’/M.9 trees
(with permission from Tustin et al., 1992).
Weeks after full bloom
2 5 8
Light regime1 Spur Bourse Spur Bourse Spur Bourse
Full sun 7.4 a2 6.3 a 7.6 a 7.0 a 7.7 a 7.9 a

Shade cloth 6.1 b 5.2 b 7.0 b 5.7 b 6.1 b 6.0 b
Natural shade 6.4 b 6.0 a 6.2 c 5.3 b 6.2 b 5.5 b
1 Light regime as follows: full sun, external canopy positions, continuously in high light conditions; shade
cloth, as full sun, but shaded to 70% shade with neutral-density shade cloth for 1 week; natural shade,
internal canopy positions, undergoing natural shading as the canopy fills in.
2 Mean separation within columns by SNK test; P = 0.05.
within 1 week. Therefore, if a bud either dif- exchanges on a leaf-area basis (specific rates),
ferentiates or develops under low photon in analogy to single-leaf determinations, it is
fluxes, it may produce leaves that inherently necessary to measure accurately the leaf area
have lower intrinsic photosynthetic poten- and the profile of photon fluxes to which
tial, with ensuing decreases in the tree’s pro- individual leaves are exposed within the
ductive potential – and the same will occur canopy. Tree leaf area is rather difficult and
under conditions that develop within a time-consuming to measure, even though
quickly closing canopy. several methods have been evaluated for its
estimation (Wünsche and Palmer, 1997a). An
alternative approach is to express tree gas
9.3.2 Whole canopy exchange in terms of the light intercepted by
the canopy (i.e. in PPF units), under the
Because different training systems have dif- assumption that most of the intercepted light
ferent light-interception profiles during the is absorbed by the tree (i.e. that reflectance
day and because orchard light interception and transmittance are negligible). This
is affected by orchard design (a combination assumption is normally valid, since leaves
of tree shape, dimensions and tree arrange- absorb, on average, 80% of the energy associ-
ment within and between rows), it is very ated with the visible spectrum.
important to study the gas exchange rates of Tree photosynthesis is a function of the
entire trees in the field over extended peri- amount of light intercepted, but it may also be
ods (Plate 9.1). However, while measure- influenced by the time of day when maximum
ment of single-leaf photosynthesis in the light interception is attained, because of the
field has become fairly easy because of the influence of temperature on photosynthetic
availability of relatively affordable commer- activity and also of other physiological deter-
cial units, at the whole-tree level progress minants. The complexity of the response is
has been slower, mostly because of a lack of demonstrated by the typical daily pattern of
commercial units, which has forced photosynthesis for a young (second leaf)
researchers to build their own instrumenta- north–south (N–S)-orientated apple hedgerow
tion – often based on adaptations of single- (Fig. 9.3): a rather sharp increase in rate in the
leaf units (Corelli Grappadelli and morning hours is followed by a slow tapering
Magnanini, 1997). off, despite the fact that light interception in
Current whole-canopy techniques pose the afternoon on the western side parallels the
specific methodological problems, which interception in the morning hours on the east-
have limited the number of studies per- ern side (L. Corelli Grappadelli and G. Costa,
formed. One of them is that, to express gas 2001, unpublished results).
3.0 8
2.5
6
Transpiration and VPD
2.0
4
NCER
1.5
2
1.0
0
0.5
0.0 –2
8 12 16 20
Time of day (h)
Fig. 9.3. Diurnal pattern of the gas exchanges of a 2-year-old ‘Gala’/M.9 apple tree, trained as a slender
spindle, in a N–S row, and of the vapour-pressure deficit (VPD), during a clear day in September. While the
carbon exchange (●) peaks in the morning hours and slowly tapers off, the water loss () increases in the
afternoon, following the pattern of the VPD (-----). Transpiration has units of mmol H2O per tree s1; VPD is
in kPa; net carbon exchange rate (NCER) has units of µmol CO2 per tree s1. (L. Corelli Grappadelli and
G. Costa, 2001, unpublished.)
Thus, if the daily pattern of light inter- total tree NCE (Giuliani et al., 1997a;
ception is different among training systems, Wünsche and Palmer, 1997b). However,
so should be the photosynthetic response. It when NCE was expressed in terms of tree
is reasonable to expect differences between leaf area, both studies showed greater spe-
the opposite sides of a palmette or central cific rates for fruiting than for non-fruiting
axis hedgerow, as opposed, for example, to trees. This has been confirmed in a more
a Y-trellis. It is known that Y-trellis orchards recent study (Wünsche et al., 2000). Removal
exhibit higher light interception than of fruit early in the season probably causes a
hedgerows or large central-leader trees. shift of resources towards greater vegetative
Row orientation will also have an impact. growth, resulting in higher tree leaf area.
While data on tree net carbon exchange This in turn may induce a greater capacity
(NCE) as a function of training systems for light interception and thus lead to the
have not been published for apple to date, higher observed NCE rates. In contrast,
similar studies in peach (Giuliani et al., fruiting trees develop less leaf area, and they
1998) have reported an increase in photo- may thus show reduced light interception
synthesis in response to increasing levels of while maintaining high specific NCE rates,
light intercepted (and have also pointed out presumably driven by the carbon demand of
that differences may exist among training growing fruit. So fruiting may not alter total
systems). It is likely that similar results tree NCE, because this appears to be a func-
would be obtained for apple, lending sup- tion of light intercepted, but it does stimu-
porting evidence to the reports of greater late the photosynthetic process, making the
productivity for those systems, such as the foliage more efficient. This has been con-
Y-trellis, that are capable of maximizing sea- firmed by chlorophyll fluorescence studies,
sonal light interception (Robinson and which have reported a decrease in photo-
Lakso, 1991; Robinson et al., 1991). chemical efficiency for the leaves of trees
Studies relating crop load and gas that had been partially or totally defruited
exchange at the tree level have initially and, as a response to this, showed a
shown a somewhat loose relationship reduced NCE per unit leaf area (Wünsche et
between the number of fruit per tree and al., 2000).
Light Relations 201
9.4 Source–Sink Relationships that heavy shade (70%) can delay by up to 2

weeks the transition of shoots from import-
The importance of different leaf types for ing to exporting.
supporting fruit growth has been widely Reduction in fruit growth rates early in
illustrated. Hansen (1971) demonstrated the the season has been correlated with subse-
dependence of fruit growth on current pho- quent fruit drop (Zucconi, 1981). Among
tosynthates, derived by spur and shoot other physiological reasons that may bring
leaves. Ferree and Palmer (1982) reported an about this reduction in growth rates, short-
important influence of spur leaves on fruit age of carbon supplied to the fruit has been
size at harvest and on fruit calcium concen- proposed as a causal agent, prior and up to
tration. Rom and Ferree (1984) reported a June drop (Schneider, 1977, 1978). Work with
strong correlation between spur leaf area and photosynthetic inhibitors (such as Terbacyl)
cumulative yields over 17 years for nine or with neutral-density shade cloth (Byers et
apple cultivars. It is important to stress that al., 1985) supports this hypothesis: inhibition
this terminology refers to the primary spur of photosynthesis up to 5 weeks AFB can
leaves, which emerge prior to the flower lead to extensive fruit drop, whereas shading
cluster and form a whorl of very efficient, at later stages does not alter crop loads
although limited, leaf area, which is capable (Byers et al., 1991) or fruit growth (Fig. 9.4).
of supporting the earliest stages of fruit Because the whorl of primary spur leaves
growth, partitioning a significant amount of is the earliest supplier of carbon to the
carbohydrates to the growing fruitlets. fruitlets, it follows that these leaves are very
The partitioning of carbon to different important in supporting the initial growth
sinks (vegetative vs. reproductive) in the tree of fruit during cytokinesis, when cell divi-
is under complex control mechanisms
(including the influence of hormones, nutri-
ents and water), but light levels can also
30
influence the directions of these flows. Early
in the season, when fruit set occurs, and
soon thereafter during cell division, fruitlets
Fruit DW (g)
receive photosynthetic carbon from primary 20 Early shade

spur leaves, which are the only carbon Late shade
source for the fruit until the developing
shoots attain sufficient leaf area to become 10
net exporters – which occurs when there are
around 13–15 unfolded leaves. Modelling
work by Johnson and Lakso (1986), subse-
0
quently confirmed by radio-tracer studies 0 100 200
(Corelli Grappadelli et al., 1994), showed the S1 S2 S3
effect of light levels on the time necessary for
the transition of a shoot from being a net
importer to being a net exporter. Shoots Fig. 9.4. Dry weight (DW) accumulation of ‘Golden
under high light levels would require com- Delicious’/M.27 fruit grown on trees that were
paratively less carbon (thus reducing their subjected to 7 days’ shading with 70% neutral-
competitive demand) and would become density shading cloth, at different times after full
bloom. Each symbol represents the average of at
exporters sooner in the season than shoots
least 40 fruits, while the lines represent the fit of the
growing in low light conditions. Short shoots
expolinear model to the actual data. One week of
(2 cm long) would begin exporting photo- shading at 5 weeks after full bloom (S1) caused
synthates sooner than long shoots (50 cm) extensive fruit drop, resulting in increased growth of
and contribute more carbon to the growing the remaining fruit (early shade). Subjecting
fruit in the early part of the season, when the adjacent trees to two similar periods of shade at
potential for final fruit size is set (Lakso et al., later stages (S2 and S3) did not cause any fruit
1989). The validation of the model revealed abscission or any change in growth rate (late shade).
sion sets the potential for subsequent fruit carbon export or reduce the flux of that
development. This probably explains the export. Studies that related the partitioning
good relationship that is normally found of light interception between primary spur
between the spur-leaf area and yield, where and bourse-shoot leaves in the early season
normally such relationships are not as good have concluded that an advantage exists for
for the long-shoot canopy (Fig. 9.5; those canopies that maximize the amount of
Sansavini and Corelli Grappadelli, 1992; spur-leaf area that is well illuminated early
Wünsche et al., 1996). The light levels avail- in the season.
able to these leaves thus become very
important, as shade can delay the onset of
9.5 The Basis of Orchard Productivity
90
(a) Theoretically, light interception sets the max-
75 imum potential for yield, as the latter cannot
60 exceed the former. From the standpoint of
light alone (but without ignoring the many
45
other factors that have an effect on yield), the
30 basis of orchard productivity lies within the
complex relationships between orchard
15
Whole canopy design components (such as tree form, tree
spacing and row orientation) and sun
(b) position. These relationships affect the two
75
essential properties required of a canopy:
concurrent capacity for high light intercep-
Yield (t h–1)
60
tion and good distribution throughout the
45
canopy. Since training-system studies are
30 expensive and time-consuming and do not
15
yield useful data before several seasons have
Spur canopy elapsed, modelling work has been instru-
mental in studying these relationships. It is
(c) appropriate therefore to start this section
75
with a brief overview of modelling work and
60 its influences on the evolution towards the
45 increased planting densities that are used in
modern training systems and orchard config-
30 urations worldwide.
15
Shoot canopy
0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 9.5.1 Models
Leaf area index (LAI)
Earlier models defined trees as solid, non-
Fig. 9.5. The relationship between fruit yield transmitting and non-reflecting canopies
(t ha1) and mean whole-canopy, spur-canopy and (Cain, 1972; Jackson and Palmer, 1972).
extension-shoot-canopy LAI (average for 3–5 and They were mostly concerned with the over-
10–12 weeks AFB) in a 4-year-old ‘Jonagold’/Mark all effect of tree dimensions and spacings
(●) and a 15-year-old ‘Marshall McIntosh’/M.9 ()
and row orientation on light interception.
slender-spindle apple-production system in 1992.
This work confirmed that the preferred row
Linear regression equations (n = 30) are: (a) yield =
12.74 + 27.37 (total LAI), r 2 = 0.656; P ≤ 0.0001; orientation should be N–S, in spite of the
(b) yield = 1.29 + 35.63 (spur LAI), r 2 = 0.774; P ≤ fact that, depending on latitude (which
0.0001; (c) yield = 41.12 + 5.75 (shoot LAI), r 2 = affects the elevation of the sun) and canopy
0.008; P ≤ 0.643. (From Wünsche et al., 1996, height, this orientation does not always
reproduced with permission.) maximize the amount of light intercepted
Light Relations 203
by a canopy. However, much more uniform (i.e. of unchanged Tc) is a preferable strat-
illumination of the canopy occurs in N–S egy, in order to increase light interception
rows, independently of latitude and time of (thus reducing Tf) without increasing
the year. poorly illuminated fractions of the canopy,
Recent studies with whole-tree chambers as would be the case if the leaf density of
on slender spindle-trained trees on N–S rows existing trees were increased by an equiva-
have revealed, however, that the diurnal gas lent amount of leaf area (Fig. 9.6).
exchanges of a tree have a complex relation- The modelling work outlining the basis
ship with light and temperature (Fig. 9.3), of orchard productivity, in terms of light
since photosynthesis normally peaks in the interception and utilization, has also been
morning hours and tapers off in the after- useful in several instances of further model-
noon, even if the amounts of light inter- ling work, as is the case in work aiming to
cepted remain high, while the loss of water estimate the daily carbon gain of an apple
via transpiration often follows the vapour- tree (Lakso, 1992). More recent work with
pressure deficit (VPD) gradient, which nor- whole-tree chambers is leading the way
mally reaches a maximum in the afternoon, towards new developments in modelling:
closely following the daily temperature results are confirming that photosynthesis at
course (Giuliani et al., 1997b). As a result, the whole-canopy level can generally be
water-use efficiency (WUE) may be quite dif- considered a linear function of instanta-
ferent at different times of the day. If further neous light interception. Subsequently, sim-
work should confirm the asymmetry of the ple and accurate methods to determine the
gas exchanges around solar noon, this might amount of light intercepted have been
pose the question of canopy orientation developed, which yield data that allow the
under an interesting new ‘light’. tracing of contour maps of light/shade lev-
Since trees do not have solid, non-trans- els on the ground under a tree (Giuliani et
mitting canopies, further refinements of al., 2000) at different times during the day.
these models took into consideration the From this information it is possible to derive
transmitting capacity of the canopy. These a three-dimensional representation of the
refinements recognized the need to ensure canopy, including an estimate of the leaf
that there are appropriate light levels area exposed perpendicularly to the sun
throughout the canopy in order to maintain (which is more closely related to transpira-
essential vital processes, such as flower- tional losses). It appears as though a reverse
bud differentiation, ensure the high photo- course might be possible where, by measur-
synthetic potential of the leaves, improve ing the amount of light intercepted, the
the tree’s carbon balance and positively physical characteristics of the canopy may
affect fruit quality traits, such as skin be able to be reproduced and these may then
colour, soluble-solids concentration and be used to estimate the tree’s gas exchanges
firmness (Plate 9.2). Jackson and Palmer (E. Magnanini, Bologna, 2000, personal com-
(1980) proposed simplified equations that munication). If the slope coefficients for the
accounted for the discontinuity in orchard linear relation between photosynthesis and
canopies. They divided incoming radiation light interception should prove to be fairly
into two components: one representing the constant (a likely hypothesis under compa-
light that is directly transmitted to the rable tree conditions), whole-tree photosyn-
orchard floor without interacting with the thesis might be accurately estimated quite
foliage (Tf); and one equal to that which simply by measuring/modelling the light
would be intercepted by the canopy if it interception of a canopy. Preliminary indica-
were non-transmitting (Fmax), minus a fac- tions, from studies including kiwi fruit,
tor accounting for transmission through the apple, peach, grape and pear whole-canopy
canopy (Tc). The latter is a function of gas exchanges (E. Magnanini and L. Corelli
planting density, tree form and canopy Grappadelli, Bologna, 2000, unpublished
density. According to this model, planting results), seem to be in good agreement with
additional trees with optimal leaf density the above assumption.
(a)
Fmax Tf Fmax –Tc Tf
(c)
(b)
Fig. 9.6. Basic concepts in light interception. (a) The theoretical maximum light interception of a solid, non-
transmitting canopy (Fmax) is reduced, in real-life orchards, by two components: Tf, the fraction of light that
misses the tree completely and is transmitted directly to the orchard floor, and Tc, the fraction of light that is
transmitted through the canopy. The former is a function of tree shape, size and spacings; the latter is a
function of the extinction properties of the foliage and the density of the canopy (the hatching reflects the
degree of shade). (b) Increasing light interception in multi-row beds via increased foliage density might
result in excessive shading of the inner canopy, while adding a row of trees without increasing canopy
density would be preferable. (c) In single-row layouts, increasing the height (but within limits dictated by the
distance between rows) will provide a similar increase in interception, without the inner canopy shading
caused by increased canopy density. (Adapted from Jackson and Middleton, 1988.)
9.5.2 Orchard configuration studies multiple rows to bed systems), similar cumu-
lative yields per hectare from the fourth to
Models have favoured orchard systems that the seventh leaf. However, rather large dif-
are characterized by narrow (or shallow) ferences existed when cumulative yields
canopies, as is the case with slender spindle, were expressed in terms of cumulative leaf
palmette (and in many hedgerow-forming area (Table 9.2). Despite different rootstocks
systems), Y-trellis and the so called ‘bed sys- (M.9 vs. M.27), a fivefold increase in planting
tems’ of short (< 2 m) trees planted very density (1800 vs. 9000 trees ha1) and widely
close together, because they can combine different orchard layouts (single vs. multiple
high light interception with very good distri- rows), the bed systems and the palmette
bution of light inside the canopy. form exhibited similar efficiency (tonnes of
Experimental evidence has confirmed that fruit per leaf area) and both were 35–45%
very different orchard designs, based on more efficient than the other systems tested,
varying tree forms, may achieve similar which were based on the slender spindle (or
yield efficiency if these constraints are satis- derived forms) in single or multiple rows.
fied. Sansavini and Corelli Grappadelli These systems had poorer light distribution
(1992) demonstrated the efficiency of narrow profiles throughout their canopies than the
canopies compared with tree forms that more efficient ones (Corelli Grappadelli and
allowed the trees to grow dense and poorly Sansavini, 1989); this difference was attribut-
illuminated canopies. They reported, across able to rootstock/training system/density
a range of tree densities, training systems combinations that were not suited to the
and orchard configurations (from single to fairly vigorous environmental conditions
Apples - Chap 09
21/3/03
2:56 pm
Table 9.2. Leaf area per tree divided between spur and shoot components and total, leaf area index (LAI), yield per hectare and fruiting efficiency (ratio of yield
to LAI) for ‘Golden Delicious’ trees trained to different tree shapes, orchard designs and densities. Leaf area, LAI and yield data are cumulative figures from the
fourth to the seventh leaf. (Adapted from Sansavini and Corelli Grappadelli, 1992).
Page 205
Leaf area per tree Fruiting
Orchard Planting (m2) Yield efficiency
layout density LAI (t ha1) (t ha1 leaves)
Light Relations
Training system (no. of rows) (trees ha1) Spur Shoot Total (a) (b) (b/a)
Slender spindle1 1 2667 6.4 c2 40.0 b 46.4 b 12.4 b 243.2 a 19.6

Slender spindle1 2 2667 6.4 c 39.7 b 46.1 b 12.3 b 218.2 a 17.7
North Holland spindle1 3 3571 5.1 b 36.0 b 41.1 b 14.7 c 253.9 a 17.3
Mini spindle3 6 8889 3.0 a 7.3 a 10.3 a 9.2 a 285.0 a 31.0
Mini bush3 6 8889 3.0 a 7.2 a 10.2 a 9.1 a 279.4 a 30.7
Palmette1 1 1802 8.7 d 36.3 b 45.0 b 8.1 a 227.6 a 28.1
1Rootstock: M.9.
2Mean separation within columns by SNK test; P = 0.05.
3Rootstock: M.27.
205
experienced in the southern Po Valley of commercial orchards (sometimes in excess

Italy. Similar differences in total yield and of 10,000 trees ha1) are all based on single-
yield efficiency in favour of systems with row layouts, with extreme reductions in
better light interception/distribution profiles the distance of the trees along the row
have been reported by other authors, com- (< 50 cm). These orchards represent extreme
paring pyramid-type canopies either with configurations, requiring highly skilled,
slender spindle and trellis systems (Ferree, extremely careful and timely management.
1989) or with the Y-trellis (Robinson and
Lakso, 1991; Robinson et al., 1991).
All these studies illustrate the impor- 9.5.3 Light distribution
tance of high light interception that was
derived by modelling work. Thus, the prac- The interception of high proportions of the
tical consequences of this work have been available PPF does not in itself guarantee
many. It has provided fundamental the production of large amounts of high-
grounding evidence for the shift towards quality apples per hectare. An even more
increased planting densities, by demon- important requirement is that the light be
strating the advantage of high-density uniformly distributed to all or most parts of
plantings of low-vigour trees in terms of the canopy. This is because all factors influ-
higher light interception early in the encing productivity are in turn influenced
orchard’s life, which resulted in earlier and by reduced light levels. The issue is then one
higher initial bearing of the trees. This, of optimizing the distribution of light within
combined with the multiplying factor of the canopy.
tree density, made possible greater initial
yields (up to 10 t marketable apples ha1 in
9.5.3.1 Orchard design
the second leaf (Sansavini et al., 1989)).
Additionally, it demonstrated the advan- From a theoretical approach, for a given
tage of smaller trees, more closely spaced, planting density, the amount of light inter-
in terms of light distribution within the cepted decreases with increasing ‘rectangu-
canopy (Jackson and Middleton, 1988), and larity’ of planting (in orchard design,
provided once again theoretical scientific rectangularity of planting is defined as the
grounds in support of the establishment of ratio of the spacing of trees between rows to
very high-density orchards, such as those their spacing along the row). However, the
reaching 6000–8000 trees ha1 (or even light distribution within the canopy may
higher) currently (Sansavini and Corelli also be negatively affected by rectangularity.
Grappadelli, 1997). One aspect suggested Working with orchard layouts designed to
by models that subsequent work has not test these hypotheses, Wagenmakers and
confirmed is related to the advantage of Callesen (1995) have reported that seasonal
growing multiple-row beds of trees, which, interception increases with decreasing rec-
contrary to the indications from modelling tangularity for a given planting density.
work, have not been performing in actual They also demonstrated that more uniform
trials as might be expected. Many reports light transmission to the orchard floor was
agree on the single-row layout as the most achieved at the highest density, with a 1 : 1
suitable for combining high productivity of rectangularity (i.e. with trees equally spaced
high-quality apples with ease of manage- between rows and along each row).
ment (multiple-row beds would require tai- However, practical considerations in general
lor-made and probably expensive prevent the utilization of such a rectangular-
equipment). The major drawback of these ity ratio, since a minimum alley width must
multiple-row plots has invariably been be maintained in order to allow room for
related to reduced light levels within the mechanical equipment to move through the
collective canopy formed by many adjacent orchard – this is one of the main reasons
trees, coupled with difficulties in manage- against the adoption of the so-called ‘bed
ment. As a result, today’s very high-density systems’ of planting.
Light Relations 207
9.5.3.2 Training systems radiation ratios (this can be the case, for
example, when comparing data from the
Based on the light-extinction characteristics
very clear, high direct radiation environment
of apple leaves, modelling work (Jackson of Washington State with those from the
and Palmer, 1980) indicates that, when leaf- mid-Atlantic states (e.g. Campbell and
area density in the outer parts of a canopy is Marini, 1992)).
increased (such as is often the case with
large, vigorously growing trees), this causes
an exponential decrease in the amount of 9.5.3.3 Pruning
light transmitted throughout the canopy. Pruning is a very important tool for guiding
This explains the very strong decrease in and controlling the growth of the tree in
light levels reported by many authors that order to maximize orchard productivity. The
have measured this parameter at different goal of pruning when the tree is young, in
positions within a tree and across different the ‘training’ stage, is to attain the desired
tree shapes (Jacyna, 1978; Tustin et al., 1988; canopy form and size in the shortest possible
Barritt et al., 1991). time; following this, the tree enters the
Since most of the plant’s characteristics mature stage and pruning goals become dif-
that influence the quality of production and ferent. In the young tree, the best approach
the overall productivity of a tree are related to maximizing light interception is often to
to light levels, it is not surprising that a great allow the plant to grow freely, without exten-
deal of effort has been put into detailing the sive removal of shoots and limbs (except for
light profiles of training systems, as well as those systems like the Y-trellis or the pal-
into making some of them more efficient by mette hedgerow that require branch selec-
targeted pruning strategies. Medium- or tion and positioning). In young trees, light
long-term studies of training systems have, distribution is normally not a concern
in general, reported similar yields of because the tree is ‘open’ to light penetra-
comparable-quality fruit from widely differ- tion, and retaining a large amount of wood
ent training systems (Ferree, 1989; Palmer et may reduce the total growth of the tree
al., 1989; Sansavini et al., 1989; Sansavini and (which is beneficial in high-density plant-
Corelli Grappadelli, 1992), provided they ings), both via an increased number of fruits
were capable of comparable, high, light and because of a lack of growth-promoting
interception. In general, however, these stud- cuts. Often in this phase of orchard life, sum-
ies have reported that fruit-quality attributes mer pruning is preferred because it is better
were more influenced than yield by light lev- suited to ‘guiding’ the tree into the desired
els within the canopy. development without leading to excessive
In addition, the type of prevailing regrowth (noting, however, that the timing
weather can have an impact on the light lev- and severity of summer pruning are critical).
els within a canopy: under climates typical When a tree reaches maturity, after it has
of the northern European countries or the filled its allotted space, the challenge
continental climate in the north-eastern USA, becomes that of maintaining flower bud dif-
which tend to have a fairly high diffuse, ferentiation throughout the canopy, as this
radiation component during the summer (for forms the basis of tree productivity. The
example, due to haze or light cloud cover), main goals of pruning a mature tree include
maximum light penetration to the interior preventing the canopy from exceeding its
canopy occurs under a mix of direct and dif- allotted space (to avoid shading neighbour-
fuse radiation, rather than under maximum ing trees) and, above all, maintaining good
direct light conditions (Lakso and light penetration through the vegetative vol-
Musselman, 1976). As a result, some of the ume, particularly for those cultivars that dif-
correlations that have been reported between ferentiate on 2- or 3-year-old wood, where it
light levels and fruit-quality parameters may becomes essential to provide adequate light
not be as close as expected if they are studied levels to maintain reproductive develop-
under climates with different direct: diffuse ment. In most modern training systems,
which try to combine low cost with high effi- that have been observed in apple, however,
ciency and ease of management, the pruning are not related to (or are not strictly con-
strategies of standard-type mature trees trolled by) phytochrome-mediated mecha-
almost always rely on branch removal (i.e. nisms. This has been demonstrated in
‘opening up’ the canopy), rather than using several studies with neutral-density shade
selective thinning cuts on existing branches, cloth, which reduces the intensity of light
in order to improve light penetration. without altering the spectral composition of
the radiation. This is not to say, however,
that none of the phenomena influenced by
9.6 Light Effects on Physiological light are insensitive to the relative composi-
Parameters tion of the spectrum.
Low PPFs in the visible range have been cor-

related with a number of negative effects in 9.6.1 Responses to light quality
apple. These include poor flower-bud differ-
entiation, reduced fruit set and shade leaf 9.6.1.1 Fruit colour
photosynthetic characteristics (such as
reduced leaf-area density, nitrogen content When fruit mature, their skin undergoes
and thickness). Fruit characters that are neg- colour formation. This is a complex phe-
atively affected by shade include fruit size, nomenon, in which, among other processes,
fruit colour, soluble-solids concentration and pigment synthesis (anthocyanins,
firmness. Also the content of mineral ele- carotenoids and flavonoids) must occur,
ments, such as calcium, and, to a lesser along with chlorophyll degradation. It has
extent, potassium, phosphorus and magne- been shown in red-fruited cultivars that
sium, in the fruit is negatively influenced by anthocyanin synthesis is under the control
the lack of light. The physiological mecha- of UV-B (< 320 nm) radiation, with a peak at
nisms behind most of these responses are not 312 nm, and that this process can be partly
fully elucidated, and the conclusion should stimulated by addition of radiation in the
not be drawn from the present discussion red region (Arakawa et al., 1985). This effect
that light is the only regulatory factor is synergistic: it causes an increase in antho-
involved, discounting other very important cyanin synthesis that exceeds the summa-
aspects, such as the influence of hormones tion of that of the two wavelengths taken
and the roles of mineral nutrition and water alone. Recent work with ‘Fuji’ trees sub-
relations. What can probably be stated is that jected to treatments with light-reflecting
light and its associated parameter, tempera- mulches (Plate 9.3) (Ju et al., 1999) has indi-
ture, are the critical environmental factors cated increased activity of a light-inducible
that trigger adaptive tree responses. The enzyme (uridine diphosphate (UDP)-
mechanisms by which these adaptive galactose:flavonoid-3-o-glucosyl transferase
responses are controlled and brought about (UFGalT)) involved in anthocyanin synthe-
are certainly many and likely to overlap in sis (Ju et al., 1995). The effect was greater for
their potential influence. This discussion will those mulches with a greater percentage of
centre only on the knowledge relative to UV and IR reflectivity. The above explains
light effects, since this is the scope of the pre- the advantage for fruit-colour formation that
sent chapter. is normally found in apples grown in alpine
As light interacts with the canopy, it regions (South Tyrol in Italy) or under very
undergoes both qualitative and quantitative clear skies in environments (such as
changes, resulting in greatly diminished Washington State and New Zealand) where
intensities and in a change in the red : far- the proportion of UV radiation is increased
red ratio towards longer wavelengths as a consequence of reduced absorption by
(Baldini and Rossi, 1987), which can trigger the atmosphere, which preferentially
phytochrome-mediated photomorphogene- absorbs more radiation in the shorter wave-
sis. Most of the light-dependent responses lengths (Nobel, 1983).
Light Relations 209
9.6.1.2 Fruiting buds that are differentiated under low light,

but also because of the lower fruiting poten-
In cultivars that have low amounts of fruit
tial of the spurs that develop under reduced
set, fruit abscission has been reduced by
light conditions. It is known that light levels
additional lighting of trees at night with red-
below 15% of full sun result in very low or
light flashes of variable duration (Greene et
absent flowering and that optimum bloom
al., 1986), an indication of the possible
occurs at light intensities above 50–60% of
involvement of a phytochrome-mediated
full sun. In addition, bloom in low light sel-
mechanism. In contrast, however, many
dom results in adequate fruit set. As fruit set
studies have indicated that fruit abscission
has been shown to depend on fruit growth
can be induced by neutral-density shading in
rate, a direct involvement of carbon
the early part of the season. Fruit abscission
resources may be assumed, but other factors
is, therefore, another example of a physiolog-
(see also Chapter 7) must be involved, for
ical process where, very probably, more than
example, in determining the direction of car-
one factor controls the responses observed.
bon flows between different sources and dif-
ferent sinks (especially vegetative vs.
reproductive sinks). However, it is known
9.6.2 Responses to light intensity
that not only do reduced light levels reduce
the rates of photosynthesis, but they also
9.6.2.1 Flower-bud differentiation
alter the patterns of carbohydrate partition-
Low light levels in the canopy result in ing among leaves. Hence low photosynthesis
reduced flower-bud differentiation. Despite and reduced hormone, nutrient and water
the fact that this observation is reported in a supply may all lead to reduced fruit set in
number of studies, the reasons for this are not shaded regions of a canopy.
fully elucidated. As floral differentiation is a On young, developing bourse-shoot
complex phenomenon, it should be expected leaves, the morphogenetic responses to
that many factors are involved, including the shade may occur within days of the imposi-
role of hormones (a promotive role for tion of shade (Table 9.1). Under natural con-
cytokinins has been proposed, while gib- ditions, therefore, these leaves may acquire
berellins inhibit flower-bud differentiation). shade leaf characteristics as, while they
How light is involved in this process is still develop, the light levels naturally and pro-
highly speculative. One hypothesis suggests gressively decrease as the canopy forms
that light might influence the direction of (normally maximum light interception is
xylem sap flow (which contains cytokinins) attained within 1 month after bloom). The
via its effect on transpiration, but this hypoth- primary spur leaves (which mature more
esis has not been conclusively confirmed by rapidly than the bourse leaves) appear, in
research work (Lakso, 1994). From a photosyn- contrast, to be less influenced by the current
thetic point of view, it is possible that reduced light environment, but they may show the
localized photosynthetic activity, due to low effects of the previous season’s light levels
PPF, may negatively affect floral differentia- under which the spur differentiated (Tustin
tion due to reduced carbohydrate fluxes. Some et al., 1992). This effect, coupled with that
evidence exists that enhancing the carbohy- affecting the developing bourse-shoot
drate status of the tree may enhance bloom in leaves, might explain in terms of photosyn-
young trees (Hansen and Grauslund, 1980, thesis the reduced levels of fruit set in
cited in Lakso, 1994) or return bloom in heav- shaded parts of a canopy.
ily cropping trees (Lakso, 1994). The duration of light periods has been
shown to influence partitioning of carbon
between different carbohydrate fractions
9.6.2.2 Fruiting
(including sorbitol, sucrose, glucose, fructose
Fruit distribution within the canopy is highly and starch (Wang et al., 1997)). The most rele-
correlated with light distribution, primarily vant fraction is sorbitol, normally represent-
because of the reduced number of flower ing more than 50% of the total carbon fixed in
apple. Short light periods (2 h) favour parti- impair the vegetative–reproductive equilib-
tioning of newly fixed carbon to sucrose (a rium of the canopy (such as weather, prun-
more readily usable carbon form in the source ing, thinning, mineral nutrition and
leaf), whereas longer light periods favour par- irrigation) can have profound effects. When
titioning of carbon into sorbitol. According to a tree carries an excessive crop, the differen-
Wang et al. (1997), sorbitol might only be syn- tiation of flower buds is reduced and this can
thesized under longer light durations and so result in a lower crop the subsequent season,
becomes an important fraction for export when intense vegetative growth can be
from the leaf (source leaves do not readily uti- expected. This vegetative flush, if uncon-
lize sorbitol, as opposed to sucrose). An addi- trolled, will result in reduced light levels,
tional element supporting this hypothesis leading to the differentiation of poor-quality
might be the fact that sorbitol synthesis spurs. This can set in motion a series of
requires nicotinamide adenine dinucleotide shade-adapted responses that can result in
phosphate (NADPH) from the light reaction loss of productivity and/or fruit quality. It is
of photosynthesis, and this might be limiting thus necessary to maintain high light levels
under short light periods. If this hypothesis throughout the canopy in order to avoid the
were confirmed, it might help explain onset of this behaviour.
reduced export from leaves that do not have Fruit from well-lit parts of the canopy tend
sufficient light energy available for the syn- to have a greater occurrence of some physio-
thesis of the export form of carbon, lending logical disorders during storage, such as bitter
supporting evidence to the role of carbon pit and internal breakdown and rotting.
shortages in shade to explain fruit abscission. Conversely, fruit from shaded parts of the
canopy show increased transpiration and
shrivelling during storage, which have been
9.6.2.3 Fruit-quality traits
linked to a greater occurrence of cracks in the
The knowledge that moderate to high light skin and with the more disorganized structure
levels are necessary to ensure adequate proof the epidermal waxes on the shaded side of
ductivity and fruit quality stems from the the fruit (Knuth and Stosser, 1987).
early determinations of light intensity, start- Illuminated fruit (or the illuminated part of a
ing from the classic work of Heinicke fruit) show greater organization of the epider-
(1966a,b) and Jackson (1967, 1968). Several mal cells, with less cracking and better struc-
authors subsequently reported 30% of avail- ture of the waxes, along with a higher fatty
able full sun as the minimum light level nec- acids content, which has been related to an
essary to ensure good-quality fruit, as well as increased capacity to control transpiration and
to maintain flower-bud differentiation (Cain, shrivelling by these fruit (Knuth et al., 1987).
1971; Lakso, 1980; Sansavini et al., 1980).
The negative effects of diminished light
levels within the canopy have been studied 9.7 Controlling Light Levels in the
in depth. Jackson et al. (1977) provided a Orchard
detailed analysis of the effects of shade on
fruit-quality parameters, including post- 9.7.1 Bagging
storage assessment of physiological dis-
orders that appear to be related to the light Following the introduction of ‘Fuji’ in the
regime under which fruit develop. Further, early 1960s (Kikuchi et al., 1997), the practice
they identified potential carry-over effects of of growing fruit in bags during development
shade from one season to the next, thus (‘bagging’) has been adopted and perfected,
introducing an additional reason for careful in order to improve fruit coloration of this
management of the light environment within often poorly coloured cultivar (Plates 9.4 and
an apple tree. The concept of carry-over 9.5). Individual fruits are placed within dou-
effects has been confirmed by other studies: ble-layered bags 4–6 weeks AFB. The bags
given the natural tendency to alternate bear- are removed in two steps 4–5 weeks before
ing of the apple tree, any cause that can the expected harvest date (Arakawa, 1998).
Light Relations 211
The outer bag, which provides the shading, is Apart from the effectiveness for colour
usually white or light-coloured, with the formation, bagging has been reported to be
internal side coloured black. The inner bag is capable of improving skin finish (Fan and
made of thinner, translucent paper, which Mattheis, 1998), reducing soluble-solids con-
can be of many different colours (red or green centration and increasing the incidence of
prevail), with the bottom open. The inner bag some postharvest disorders, including bitter
is removed a few days (up to 1 week) after pit (Witney et al., 1991) and scald (Fan and
the outer, and can contain waxes and fungi- Mattheis, 1998). Despite its positive effects,
cides to protect the fruit from bruising and bagging individual fruit in the orchard is
diseases. In Japan, this technique has been expensive and needs to be evaluated from an
extended to other cultivars, including economic standpoint. Indications from Japan
‘Mutsu’ (a non-red apple), and has been and the Pacific north-west region of the USA
tested and adopted in several apple-growing are that the use of bagging is declining
areas of the world, with adjustments to adapt because of cost, while, for the same reasons,
it to different light and temperature condi- it has never been adopted on a commercial
tions (Fan and Mattheis, 1998). basis in other important apple-growing areas
Coloration of red apples is due to the syn- (Italy, for example).
thesis of anthocyanins, which depends pri-
marily on light intensity and quality
(Arakawa et al., 1985), temperature 9.7.2 Reflectants
(Arakawa, 1991) and many other factors, as
reviewed by Saure (1990). Along with the The knowledge that light can influence many
synthesis of anthocyanin, it is very important vegetative and reproductive phenomena has
that chlorophyll degradation occurs, which led to investigations that have evaluated the
also takes place during the final stages of possibility of increasing the light available in
fruit ripening. Reducing the amount of the lower and inner parts of the canopy
chlorophyll synthesized by placing the fruit through the use of reflecting materials (also
within a non-transmitting bag may help in termed reflective mulches). Such increases in
improving skin coloration at harvest pro- light availability might have beneficial
vided the fruit is re-exposed before harvest effects on tree performance. Most, but not all,
in order to allow the light-dependent pig- of the materials tested include aluminium as
ment synthesis to occur. Proctor and a reflecting agent, coated or otherwise
Lougheed (1976) reported decreased chloro- applied to different types of supporting
phyll content and increased colour formation material, including tar-paper, polypropylene,
in apples that had been placed in aluminium polyethylene or polyester films. White, non-
bags from about 4 weeks AFB and exposed metallized polypropylene film can also be
20–30 days before harvest. used. The main conclusions from work with
These observations and the underlying these materials include the following.
mechanism have been confirmed by other
studies. Arakawa et al. (1985) showed the
9.7.2.1 Light levels
dependence of anthocyanin formation on
UV-B (< 320 nm) light. They also indicated Reflectants do improve the light levels in the
that different cultivars have different tem- lower, inner parts of the canopy (Doud and
perature optima for colour formation. Ferree, 1980b; Mika, 1980), although their
Cultivars that do not colour well (e.g. ‘Fuji’) effectiveness decreases with the ageing of the
exhibit low temperature optima, between 15 material and it may also vary depending on
and 20°C, and this condition needs to be cou- the type of natural light environment. Mika
pled with high UV availability. The timing of (1980) reported a decrease of about 50% in
bag removal is fairly critical for colour for- reflectivity of aluminized tar-paper within 3
mation, as fruit seem to acquire and then years of its deployment in the orchard. Doud
lose the capacity for anthocyanin synthesis and Ferree (1980b) reported a greater increase
with time, as maturity approaches. in light penetration within the canopy under
cloudy conditions than during sunny days. cyanin synthesis and of chlorophyll break-
This may be related to a greater ‘light-har- down. They reported increased ethylene lev-
vesting’ capacity of some of the reflective els due to the reflectants, as well as increased
materials under diffuse light conditions. activity of a key enzyme in the biosynthetic
pathway of anthocyanin formation – UFGalT
– an enzyme that is light-inducible. The
9.7.2.2 Fruit size and colour
increased anthocyanin synthesis recorded in
An increase in fruit size in the lower canopy parts of the trees with increased light
of apple trees was reported as a consequence because of the reflection might be due to a
of use of reflectants between 50 and 100 days stimulatory effect of light on gene expres-
before harvest (Moreshet et al., 1975), contrary sion, with a possible involvement of ethylene
to the reports by Doud and Ferree (1980a), in the regulation of such expression. This
Mika (1980) and Ju et al. (1999). However, all method too, however, requires careful eco-
these studies agree on increased apple skin nomic evaluation, since its costs are not neg-
colour in the fruit from the lower, inner parts ligible, in particular those related to its
of the canopy as compared with treatments disposal (the reflectants used normally
where no reflective mulch had been used. remain in the field for more than one season,
until they lose reflectivity or become too
damaged to remain effective).
9.7.2.3 Physiological processes
While the early studies did not indicate any
effects of the reflectants on photosynthesis or 9.7.3 Shading to reduce sunburn
transpiration, a recent detailed study of the
absorption of a tree under which a reflective Apple fruit exposed to high light intensities
ground cover had been applied (Green et al., during the growing season may develop sun-
1995), while confirming an increase in the burn damage, which is related to excessive
light available inside the lower canopy and temperatures in the fruit surface tissues. The
subsequent greater absorption of energy by intensity of damage can vary, but in all cases
the tree, also reported increased transpira- the commercial value of the fruit is lost or
tion due to the greater radiation load on the greatly reduced. Fruit temperature may vary
leaves and coincident higher photosynthesis between the exposed and the shaded fruit
than from the same tree without a reflectant surfaces by more than 20°C, and the fruit
ground cover. These authors reported reacts to this variation with the synthesis of
increases of 40% in total net all-wave radia- heat-shock proteins, which impart protection
tion when the full alleyway was covered from extreme temperature to these tissues.
with the reflectant and this increase was However, under very high light levels and
accompanied by increased net photosyn- warm, dry conditions, the extent of sunburn
thesis (+34%) and transpiration (+26%). may be quite high. As a remedy, shading nets
More recently, with the widespread pro- are used to cover the orchard. This is not only
duction of cultivars such as ‘Gala’ and ‘Gala’ expensive, but may have negative effects on
strains or ‘Fuji’, reflectants have received tree productivity (Widmer, 1997). Normally
renewed attention as a management tool to white (i.e. highly reflecting) nets are used, or
improve skin coloration. This approach is evaporative cooling may be used for the
alternative to bagging, and it aims to reap same reason (however, this may also prove
the same benefits without incurring the high quite expensive where water availability is
costs of that technique. In this case, the reduced). Recently, applications of white
reflective mulch is applied 2–3 months materials for coating the foliage and fruits
before harvest (Andris et al., 1998). Recent have been proposed, based on kaolin clay
work (Ju et al., 1999) has shown that materi- particles, which aim to reflect the light.
als with high reflectivity in the UV region Experimental work is still under way for this
can be quite effective in promoting red latter technique, which, if viable, would have
colour formation, via a stimulation of antho- the advantage of low cost.
Light Relations 213
9.8 Conclusions the south-eastern Po Valley. Proper dormant

and summer pruning strategies must be
It can be concluded from this discussion that implemented to maintain the high levels of
the first and foremost concern for the apple light penetration into a canopy. This is
grower must be to ensure high levels of light essential for flower bud formation through-
interception in the orchard, coupled with out the canopy and to ensure that the buds
homogeneous distribution of light through- are of high fruiting potential. Along with
out the tree canopy. This is achieved by N–S- good flower formation and fruit set, high
orientated rows, with trees trained to forms light levels inside the canopy also promote
that have a high area/volume ratio, i.e. the formation of leaves with good photosyn-
which are thin and have a high proportion thetic potential and allow fruit to attain
of the foliage exposed to the incoming radia- large size and good eating qualities. Recent
tion. The choice of training system, however, developments that provide the possibility of
must be made keeping in mind the condi- physically manipulating light in the orchard
tions that determine the growing potential (reflectants, shading nets, kaolin-based
of a site. For example, while it is possible to sprays) might be setting the scene for an
grow double- or triple-row beds under low- array of new tools by which the growth and
vigour conditions, such as occur in alpine development of the trees may be manipu-
environments, it would be impossible to lated, simply by affecting the most impor-
adopt the same orchard design under vigor- tant environmental signal to which trees are
ous conditions, such as those occurring in sensitive: light.
References
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var, stage of fruit ripening and bagging. Journal of Horticultural Science 66, 763–768.
Arakawa, O. (1998) Colouring of ‘Fuji’ apples by bagging. Compact Fruit Tree 31, 34–35.
Arakawa, O., Hori, Y. and Ogata, R. (1985) Relative effectiveness and interaction of ultraviolet-B, red and
blue light in anthocyanin synthesis of apple fruit. Physiologia Plantarum 64, 323–327.
Baldini, E. and Rossi, F. (1987) Radiant energy spectral distribution in fruit tree canopies. In: Prodi, F.,
Rossi, F. and Cristoferi, G. (eds) Agrometeorology. Editrice Compositori, Bologna, pp. 401–408.
Barritt, B.H., Rom, C.R., Konishi, B.J. and Dilley, M.A. (1991) Light level influences spur quality and canopy
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10 Temperature
John W. Palmer,1 Jean P. Privé2 and D. Stuart Tustin3

1The Horticulture and Food Research Institute of New Zealand Ltd, Nelson Research
Centre, Motueka, New Zealand; 2Agriculture and Agri-Food Canada, Boutouche, New
Brunswick, Canada; 3The Horticulture and Food Research Institute of New Zealand
Ltd, Hawke’s Bay Research Centre, Havelock North, New Zealand
10.1 Latitude 217

10.2 Freeze Injury 218
10.3 Dormancy 224
10.4 Spring Frosts 225
10.5 Spring Temperatures 227
10.6 Length of Growing Season 228
10.7 Assimilation and Respiration 228
10.8 Fruit Growth 229
10.9 High Summer Temperatures 230
10.10 Root Growth 230
10.11 Shoot and Leaf Growth 231
10.12 Synthesis of Temperature Effects in Models 231
10.13 Fruit Quality 232
Temperature has profound effects on all 10.1 Latitude

aspects of apple production. First, it sets
boundaries on production areas. Being basi- The genetic pool of the cultivated apple has
cally a temperate deciduous species, the culti- resulted in a very adaptable species. Large-
vated apple needs a period of winter chill to scale commercial apple production is gener-
break dormancy. Secondly, temperature con- ally limited to the latitude range of 25 to
trols the length of the growing season, which 52°. Apples can be grown outside this range
in turn limits the range of cultivars that can be if conditions are favourably warm in higher
grown in any one location. Thirdly, tempera- latitudes by virtue of being close to a water
ture alters the rate of development of all phys- mass (e.g. Norway 60°N) or favourably cool
iological processes, including key processes in lower latitudes by virtue of elevation.
such as the rates of pollen tube growth, cell Alternatively, at low latitudes the breaking
division and respiration. Fourthly, tempera- of dormancy can be assisted by the use of
ture alters the development of apple pests and specific chemical sprays, deleafing or the
diseases so that warmer areas frequently have use of low-chill-requirement cultivars.
more generations of pests than cooler ones. Although apples can be grown in regions

218 J.W. Palmer et al.
where winter temperatures drop to 40°C, 10.2 Freeze Injury

it is frequently in practice the rootstock that
is killed by low temperatures rather than Frost injury to flowers in the spring or freez-
the scion cultivar. ing injury in the autumn or winter, which
Temperature at a site is approximately kills tissues or trees, accounts for greater
determined by latitude. Charles-Edwards crop losses than all other environmental
(1982) suggested, from a relatively small stresses combined (Flore and Howell, 1987).
data set, that average mean temperature It has also been estimated that the economic
decreases by 0.45°C per degree of latitude loss from freeze damage and cold injury to
from 10 to 55°, while the relative seasonal the USA’s fruit industry exceeds the losses
amplitude in temperature increases by from insects, diseases, rodents and weeds
0.015°C per degree of latitude over the combined (Larsen, 1970). Although several
same range. Due to the large quantity of trends in cultivation have led to a reduction
ocean in the southern hemisphere, temper- in the impact of winter freezes over the years
atures for the same latitude tend to be – namely site selection, better management
somewhat cooler in the southern hemi- practices and the use of hardy cultivars and
sphere than in the northern hemisphere. rootstocks – the ubiquitous and unpre-
For example, mean maximum and mini- dictable nature of low-temperature cycles
mum temperatures in Hawke’s Bay, New continues to cause extensive losses to the
Zealand, at 39° 40’S are cooler than Davis, apple industry. Examples include the
California, at a similar latitude of 38° 32’N 1986/87 freeze in Poland, which killed over
13 million trees (25–30% of the total apple
(Table 10.1). In contrast, when comparisons
acreage), the devastating freeze of 1984/85 in
are made between temperatures in New
western Europe (many millions of trees
Zealand and those in Japan at a similar lati-
killed, actual numbers not determined) or
tude, New Zealand temperatures are cooler
the November 1955 and December 1964
in the summer and warmer in the winter
freezes in the Pacific north-west of North
(Table 10.1). Although Aomori is at a simi-
America, which killed 2.5 million fruit-trees
lar latitude to California, the presence of
and injured 4 million trees in New York
the large land mass to the west of Japan
State, respectively. Although of less magni-
results in considerably cooler winter tem-
tude, it is common in the colder apple-
peratures than those that occur in Davis,
growing regions for substantial tree-fruit
where prevailing winds come off the Pacific losses to occur relatively frequently. For
Ocean and there is also a strong influence example, severe freezes that kill apple trees
of the Sacramento River. The amelioration or lower yields occur every 5–7 years in east-
of climatic extremes when comparing sites ern and western Canada (Coleman, 1992;
with a maritime or continental climate at a Hall and Quamme, 1994).
similar latitude in the northern hemisphere Apple trees, like other hardwood trees,
is seen in Table 10.2, where East Malling, begin to harden in the autumn from the outer
Kent (lat. 51° 18N, long. 0° 26E), is com- shoots down the trunk to the ground line, so
pared with Skierniewice, Poland (lat. 51° injury is often observed on the trunks rather
58N, long. 20° 10E). The maritime environ- than on the shoot tips. Concurrently, not all
ment of Kent does not experience the range tissues within the same genotype have the
of winter or summer temperatures experi- same hardiness (Cain and Anderson, 1976).
enced at Skierniewice. Even within a small Overall, fruit buds are more sensitive than
area, there can be considerable changes in shoots, and roots are more sensitive than
temperature due to altitude, slope and shoots, with the cambial tissue being more
slope orientation and closeness to large sensitive than the xylem tissue. Under natural
bodies of water. In the extreme case of conditions, twigs can survive to 38°C but
altering temperature with altitude, apples even the hardiest rootstocks can only survive
can be grown close to the equator at high to 18°C (Potapov, 1999). Although roots are
elevations in countries such as Ecuador. consistently less hardy than stems, this is
Apples - Chap 10
Table 10.1. Comparison of mean minimum and maximum temperatures (°C) at three sites at similar latitudes – Davis, California, USA, lat. 38° 32N, long. 121°
46E, Havelock North, New Zealand, lat. 39° 40S, long. 176° 53E, and Aomori, Japan, lat. 40° 49N, long. 140° 46E. (Temperatures for New Zealand have
been reversed by 6 months to make comparisons between hemispheres easier.)
21/3/03
Mean
Location temp. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.
Davis, USA Min. 7.8 2.3 4.1 5.3 6.7 9.5 12.3 12.9 12.6 11.5 8.7 5.1 2.5
Havelock North, Min. 7.0 1.8 3.2 4.8 6.5 8.4 10.7 12.0 12.0 10.3 7.2 4.4 2.1
2:56 pm
New Zealand
Aomori, Min. 6.6 3.9 4.0 1.4 3.8 8.8 13.8 17.8 19.7 14.9 8.2 2.8 1.1
Japan
Davis, USA Max. 23.0 11.5 15.5 17.9 21.8 26.6 31.0 33.7 33.0 30.5 25.6 17.3 11.6
Page 219
Havelock North, Max. 18.8 13.4 14.3 16.4 18.7 20.7 22.2 23.9 23.8 22.2 19.6 16.6 14.1
New Zealand
Aomori, Max. 13.5 0.7 1.0 4.9 12.1 17.4 20.8 24.4 26.8 22.8 16.8 9.9 3.8
Japan
Temperature
Table 10.2. Effect of maritime versus continental climate at two sites in northern Europe (East Malling, Kent, UK, lat. 51° 18N, long. 0° 26E, and Skierniewice,
Poland, lat. 51° 58N, long. 20° 10E) on mean monthly maximum and minimum temperatures (°C).
Mean
Location temp. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.
East Malling, Min. 5.6 0.8 0.8 2.1 4.0 6.5 9.5 11.6 11.3 9.4 6.3 3.4 1.8
UK
Skierniewice, Min. 4.3 4.4 3.3 1.2 3.3 8.0 11.3 12.9 12.5 9.2 5.0 0.6 2.7
Poland
East Malling Max. 14.0 6.7 7.0 9.9 12.9 16.5 19.9 21.7 21.5 19.0 14.8 10.2 7.6
UK
Skierniewice, Max. 12.6 1.2 2.4 6.2 13.0 19.2 21.8 23.7 25.2 18.3 12.8 5.4 1.9
219
Poland
closely associated with soil temperature, since tree survival. Root-injured trees leaf out in
stems below ground are no more resistant to the spring, but new growth and blooms are
cold than roots, and exposed roots can be as retarded and often wilt after hot weather. At
hardy as above-ground stems. There also this time, the scion often does not show signs
exists evidence that rootstocks may impart of the oxidative browning that is typical of
hardiness to the trunks and that reciprocal direct freezing. Root injury is often seen to
effects also exist for the scion (Embree and occur in the upper soil levels, with the crown
McRae, 1991). Normally, apple production and roots below a critical soil depth often
can extend into regions experiencing winter uninjured. Although most surviving roots
minimum temperatures in the range between that have vascular connections with the
34 and 40°C because the low-temperature scion will probably produce new roots, even
exotherm of apple xylem (37 to 39°C) falls a small amount of root injury can reduce
within this temperature range. When damage yields in subsequent years. None the less, an
to the xylem occurs, it is as ‘blackheart’, a con- apple tree can lose a considerable amount of
dition involving the darkening of the xylem. its root system and still survive if the crown
Repeated injury to the xylem weakens the remains healthy.
tree, promotes the invasion of wood-rotting Freezing injury occurs in many forms and
organisms and can lead eventually to death at different times of the year – it is not a sin-
(Quamme et al., 1982). However, even with gle problem. Although most research has
50% blackheart injury, it is possible for a tree focused on midwinter hardiness, the
to recover (Warmund et al., 1996). Other visi- acclimatization and deacclimatization char-
ble symptoms of freezing injury include stem acteristics of many cultivars and rootstocks
dieback, winter kill of dormant vegetative or may predispose them to injury from low
flower buds, frost splitting of tree-trunks (also temperatures in the autumn and early
known as winter sun-scald or south-west spring. The most important factor affecting
injury), crown and root injury and soil-heav- the occurrence and severity of winter injury
ing damage. Freeze damage to shoot tips and in Finland and in New Brunswick, Canada,
vegetative and/or reproductive buds is com- was mild weather during midwinter fol-
mon but affects mostly the current season’s lowed by low temperatures (Kaukovirta and
crop. South-west injury or splitting of the Syri, 1985; Coleman, 1992). Concurrently, the
bark on the south-west side of the tree is also capacity of apple roots to retain cold-
quite common in the northern hemisphere hardiness during intermittent short periods
and occurs on cold days during the winter of warm weather, i.e. freeze–thaw cycling, is
when the sun increases the temperature of the also very detrimental to survival (Privé et al.,
south-west side of the tree while it is frozen 2001). Two freeze–thaw cycles at 3°C were
(Kesner and Hansen, 1976). Temperature dif- as detrimental to root hardiness as one cycle
ferentials of >16.7°C can exist between the at 9°C. Forsline (1983) also found that a ‘so-
cambium on the south and north side of the called hardy’ rootstock, ‘Robusta 5’, lost
trunk when ambient temperatures are below more than 6.7°C of hardiness after the first
0°C. When the elevated bark temperatures are winter thaw. In response to all these different
followed by subsequent freezing, the cortical types of freezing injury, researchers have
tissues are killed and the trunk cracks. developed methods to identify the environ-
Although serious, the tree usually recovers mental variables most pernicious to produc-
after 2–3 years if secondary pathogens have tion. For example, in the Okanagan valley of
not further weakened the tree. Reflective British Columbia, Canada, the main climatic
paints and tree wraps applied to tree-trunks factor associated with poor production the
can help provide several degrees of protection next year was the adverse impact of low
for the tree and reduce or even eliminate temperature (8.5°C) during late October
south-west injury (Sakai, 1966). The most and early November (Caprio and Quamme,
damaging freeze injury to the tree occurs 1999). This information helps breeders and
when the crown and roots are affected, with growers alike to develop strategies best
the crown being the most critical to whole- adapted for their region.
Temperature 221
The basis for cold resistance is in the severe winters. Through the efforts of breed-
genetic constitution of the plant, so breeding is ers, new successful hardy introductions
the preferred long-term approach to survival. include ‘Fantazia’, ‘Fireside’, ‘Haralson’,
This has been the central objective of numer- ‘Honeycrisp’, ‘Lobo’, ‘Katja’, ‘Mantet’ and
ous breeders across Canada, the northern ‘Northern Lights’ (Zagaja, 1994). Winter-hardy
USA, Sweden, Poland, northern China and rootstocks derived from northern breeding
Russia (Zagaja, 1994). Malus baccata L., Malus programmes include Alnarp 2, Bemali, the
prunifolia Willd. and selections from Malus Budagovski series from Russia, Mark, Ottawa
sieversii Ldb. were used to transfer the poly- 3, the Kentville select clones from Canada and
genic traits of acclimatization, cold tolerance the P-series from Poland (Webster, 1999).
and avoidance to their new selections Cold-hardiness screening is also being done
(Korban, 1986). M. baccata L. was especially for the fire-blight-tolerant rootstocks from the
useful in the development of cold-hardy root- Cornell/Geneva breeding programme in the
stocks (Stepanov, 1974). Half a century ago, USA and the Vineland series from Canada
the hardy cultivars available for marginal (J.-P. Privé, personal communication). The
areas included ‘Duchess of Oldenburg’, lowest survival temperatures for some apple
‘Yellow Transparent’, ‘Wealthy’, ‘Charlamoff ’ cultivars and rootstocks between November
and several ‘Antonovka’ types, all of relatively and February in British Columbia, Canada, are
poor fruit quality but capable of surviving listed in Table 10.3, while general cold-
Table 10.3. The hardiness of 13 apple cultivars and seven rootstocks expressed as the lowest survival
temperature (°C) (from Chilton et al., 1994; Quamme et al., 1999).
Date
November December February Mean
Cultivar
‘Mutsu’ 27.9 32.9 24.5 28.5 a
‘Jonagold’ 28.4 31.2 25.9 28.5 a
‘Golden Delicious’ 26.0 33.0 26.5 28.5 a
‘Gala’ 29.0 35.6 30.3 31.6 b
‘Sunrise’ 30.9 35.8 28.8 31.8 b
‘Elstar’ 31.4 35.4 28.3 32.2 bc
‘Granny Smith’ 31.7 34.6 30.8 32.4 bc
‘Red Rome’ 32.4 35.7 32.1 33.4 bcd
‘Braeburn’ 34.0 35.7 30.6 33.5 bcd
‘McIntosh’ 33.0 37.2 34.7 35.0 cde
‘Spartan’ 35.4 37.6 32.6 35.2 de
‘Fuji’ 35.9 37.7 33.8 35.8 de
‘Empire’ 36.1 38.4 35.9 36.8 e
Mean 31.8 a 35.5 b 30.6 c
Rootstock
M.7 – 7.7 7.2 7.5 a
M.9 – 10.0 9.2 9.6 b
A.2 – 11.2 11.2 11.2 c
J.9 – 9.4 12.3 11.8 c
B.9 – 12.2 12.5 12.3 cd
O.3 – 12.0 14.2 13.2 de
P.2 – 12.6 15.7 14.2 e
Mean – 11.0 a 11.8 b
Numbers in column and row followed by the same letter are not significantly different (P = 0.05)
according to Duncan’s multiple-range test.
hardiness classifications are provided in Tables probably due to this rootstock’s poor ability to
10.4 and 10.5. Besides their overall capacity to regenerate new roots.
tolerate or avoid cold stress, there also exists At extremely low temperatures, apple tis-
some genetic variability in the ability to sues have evolved mechanisms to either
recover from a damaging cold event. The root avoid or tolerate the ice formation within
system of certain rootstocks , such as Beautiful their tissues, which causes cell death.
Arcade, B.118, B.490, and M.26, seem better Generally, xylem tissues avoid freezing
adapted to recovery than M.9, M.7, MM.111 or injury by freezing-point depression or deep
O.3 (Privé and Embree, 1997). Interestingly, supercooling, while bark and buds tolerate
some rootstocks, such as O.3, have good mid- low temperatures by preventing intracellu-
winter hardiness but poor recovery. This is lar ice formation (Quamme et al., 1982; Kang
Table 10.4. Classification of the winter-hardiness of apple cultivars based on reported differential cold-
hardiness after test winters and artificial freezing tests (from Forsline, 1983; Chilton et al., 1994;
Quamme et al., 1999; Friedrich and Fischer, 2000; A. Czynczyk, personal communication).
Tender Moderately tender Hardy Very hardy
‘Cox’s Orange Pippin’ ‘Braeburn’ ‘Alwa’ ‘Antonovka’

‘Golden Delicious’ ‘Delikates’ ‘Cortland’ ‘Columbia’
‘Gravenstein’ ‘Elstar’ ‘Empire’ ‘Dolgo’
‘Jonagold’ ‘Gala’ ‘Fantazia’ ‘Haralson’
‘Jonathan’ ‘Granny Smith’ ‘Fuji’ ‘Kerr’
‘Melrose’ ‘Ligol’ ‘HoneyCrisp’ ‘Mantet’
‘Mutsu (Crispin)’ ‘Northern Spy’ ‘James Grieve’ ‘Norland’
‘Newtown’ ‘Red Boskoop’ ‘Jersey Mac’ ‘Rescue’
‘Ontario’ ‘Red Delicious’ ‘Lobo’ ‘Wealthy’
‘Sinta’ ‘Redkroft’ ‘Lodel’
‘Red Rome’ ‘McIntosh’
‘Sawa’ ‘Melba’
‘Sunrise’ ‘Northern Lights’
‘Spartan’
‘Vista Bella’
Table 10.5. Classification of the winter-hardiness of apple rootstocks based on reported differential cold-
hardiness after test winters and artificial freezing tests (from Privé and Embree, 1997; Potapov, 1999;
Quamme et al., 1999; Zhabrovsky and Samus, 1999; Friedrich and Fischer, 2000; A. Czynczyk, personal
communication).
Very tender Tender Moderately hardy Hardy Very hardy
M.7 J-TE-D B.118 Antonovka Beautiful Arcade

Malus halliana J-TE-E B.490 Alnarp2 Malus baccata
MM.104 J-TE-F J.9 B.9 Columbia
M.2 MAC.1 KSC 7 Robusta 5
M.4 MM.106 KSC 14
M.9 MM.111 KSC 28
M.11 O.3 M.26
P.16 P.1 P.2
P.59 P.14
P.60 P.18
Supporter 4 (Pi-80) P.22
Y.P.
Temperature 223
et al., 1998). Tissues exhibiting deep super- involves both an increase in tolerance to
cooling can maintain supercooled cellular dehydration and an increase in the amount
water to 38°C and involve freeze avoid- of unfreezable water (Kang et al., 1998).
ance to the point of spontaneous nucleation Freezing injury can occur even when tis-
of the cellular solutions (Burke et al., 1976). It sues are fully hardened, because the dehy-
is this degree to which deep supercooling dration/rehydration cycles involved in
occurs that determines the higher latitude freezing have dramatic effects on both mem-
limits of apple production. However, under branes and their cellular ultrastructure. The
certain circumstances, certain cells can retain thermal stress placed on membranes during
cellular water in a supercooled state to tem- cooling can result in transitions from the
peratures as low as 47°C. Lindén et al. liquid crystalline to the gel phase, which has
(1996) reported LT50 of 46° and 43°C, important deleterious consequences. As a
respectively, for xylem tissue of the apple result of the transition, the cell may leak its
cultivars ‘Antonovka’ and ‘Samo’ grown in contents to the surrounding medium, possi-
southern Finland. Interestingly, dehardening bly due to defects in the bilayer between gel
treatments at 14°C decreased this resis- and liquid crystalline domains. In addition,
tance by 12–15°C. Compounds that help tis- the transition may lead to phase separation
sues supercool and help prevent or of membrane constituents in the plane of the
minimize injury during freezing are known membrane (Quinn, 1985). Seasonal ultra-
as cryoprotectants and include sugars, structural changes are also involved in the
betaines, amino acids, polyols, oligosaccha- changes in the physical and functional prop-
rides, antioxidants and antifreeze proteins, erties of cold-adapted apple cells. The ability
such as dehydrins (Kaye and Guy, 1995). In of cells to adjust the balance of metabolites
bark tissues and buds, ice first forms extra- between vacuoles and the cytoplasm
cellularly, and then intracellular water reversibly and to synthesize new compo-
migrates to the extracellular ice crystals due nents within the cytoplasm can influence the
to the existing water-potential gradient tissue’s resistance or susceptibility to the
(Sakai, 1965). Ice does not penetrate the cell stress-producing effects of the dehydration
membrane and therefore intracellular freez- that occurs during freezing (Kuroda and
ing is avoided. With proper treatment, buds Sagisaka, 1993).
from most apple cultivars can even survive Although breeding is the preferred
exposure to liquid nitrogen temperatures of approach to survival, this takes time, so cul-
196°C, and this is the preferred method for tural practices that help avoid or tolerate
germplasm preservation (Tyler and freezing injury are often used in marginal
Stushnoff, 1988). Sakai (1965), however, areas of production. Usually, any practice
could not achieve survival of xylem ray that helps maintain a healthy tree will be
parenchyma in liquid nitrogen. In both veg- beneficial for survival, so tree condition is as
etative and floral buds of apple, the ice crys- important as its genetics. Cold acclimatiza-
tals are formed mostly in the scales and axis, tion is an active process and requires energy
but some crystals can also be found in the and a carbon source, so it is not surprising
bud apices or flowers. This is unlike peach, that a certain minimum level of sugars or
which does not form ice crystals directly in carbohydrates is necessary for the develop-
the floral parts. Therefore, injury in peach ment of maximum winter-hardiness. Thus, it
occurs only from excessive dehydration is also expected that shading, excess crop-
rather than from freezing per se (Vertucci ping, foliage damage, poor nutrition, very
and Stushnoff, 1992). Although the amount dry or wet soils and heavy pruning can
of water in plant cells is negatively corre- reduce the trees’ ability to tolerate freezing
lated to cold-hardiness, it is the fraction of temperatures (Flore and Howell, 1987).
unfrozen water within the cells that is Early natural defoliation is believed by
important for survival (Bittenbender and growers to be an indicator of early acclimati-
Howell, 1975). Thus, it appears that the tol- zation to winter damage, i.e. the earlier the
erance of apple buds to freezing injury leaf fall, the greater the acclimatization. This
view is supported by evidence that actively growth following defoliation is used to pre-
growing shoots are less hardy than those vent the tree entering dormancy in some
that have stopped growing. However, ear- areas of the subtropics, where there is inade-
lier leaf fall is not always associated with quate chilling; in this context the trees will
greater hardiness. Earlier leaf abscission flower after defoliation and the trees can
results in fewer total carbohydrates being bear fruit twice each year. In temperate
produced by the leaves, and this may reduce regions, however, the buds become dormant
hardiness for trees that are low in vigour or and will not begin growth until a period of
for trees that have cropped heavily (Flore winter chill is followed by warmer condi-
and Howell, 1987). Likewise, nutrition and tions in the following spring. This is a com-
fertilization play a role in cold tolerance, but mon occurrence among temperate tree
it is most often overplayed. Generally, trees species and enables the tree to withstand the
that are adequately supplied with nutrients cold conditions of winter. Winter dormancy
are hardier than those that are undersup- is also known as rest, true dormancy or
plied or have excessive nutrients available. endodormancy (Lang, 1989). As temperature
Although the actual mode of action of nitro- is the main environmental factor controlling
gen on cold-hardiness has not yet been dormancy, several attempts have been made
determined, it is likely to be via carbohy- to quantify the relationship between chilling
drate accumulation. Recent studies suggest and temperature. The earliest attempts were
that postharvest foliar application of nitro- made based on the number of hours below
gen can be beneficial to hardiness by the 7°C. Most commonly used today is the non-
remobilization of this nitrogen (mostly from linear temperature-response Utah model,
rubisco) out of the leaves and into the buds, where dormancy is satisfied by the accumu-
wood and roots before leaf fall. This prac- lation of a set number of chilling hours
tice, however, may cause the leaves to (Anderson and Seeley, 1993), or one of a sim-
remain physiologically active longer and ilar form described by Shaltout and Unrath
might affect leaf fall, depending on location. (1983). In the latter model the temperature
Tree losses from root injury are often found response is a parabolic one, with maximum
on sandy or gravelly soils because these chilling efficacy at 7.2°C and zero responses
soils retain less moisture and consequently below 1.1°C and at 15°C.
have reduced heat capacity. However, wet Temperatures higher than 15°C produce
soils can have lower temperatures because an increasingly negative effect on chilling,
of greater heat conductivity, so the best soils i.e. a negation of previous chilling effects
for reducing freezing injury are those where (Fig. 10.1). Cultivars vary widely in their
conductivity is low and heat capacity is chilling requirement, with, for example, 218
high. Vegetative ground covers and/or units required for ‘Anna’ through to 1516
mulches, combined with a trapped insulat- units being required for ‘Wright No. 1’
ing blanket of snow, may be the most effi- (Hauagge and Cummins, 1991), although
cient and expedient means of preventing the majority of cultivars are in the 800–1200
winter injury to the roots from low soil tem- range of chill units. This wide range of chill-
peratures (Privé, 1994). ing requirements enables apples to be grown
from the subtropics in Israel, Brazil and
Mexico to higher latitudes such as Canada,
10.3 Dormancy Norway or Russia. The negation of winter
chill by high temperatures is a particular
Other than on very young trees, vegetative problem in warmer climates, such as those
growth of apples is complete after about 2–3 that occur in Israel and South Africa. The
months, with the formation of terminal rest- model of Erez et al. (1990) was developed for
ing buds. Although these buds can be these warmer conditions. Although warmer
induced to grow again in the current season conditions can negate chilling, experimental
by defoliation, they lose this ability as the work suggested that this was only a partial
trees pass into autumn. This ability to start negation, as the negation depended upon
Temperature 225
1.0 hours (GDH) above a base temperature of

0.8
0.6
4.5°C (Hamer, 1980), so that about 290 GDH
0.4 are required for full bloom of both
0.2 ‘Delicious’ and ‘Cox’s Orange Pippin’.
0.0
Hamer assumed a linear response to temper-
Chill units
–0.2
–0.4 ature above the base temperature. This
–0.6
–0.8
assumption is frequently used in models
–1.0 based on thermal time, as is the assumption
–1.2 of a constant base temperature. Other work-
–1.4
–1.6 ers, e.g. Winter (1986) and Anderson and
–1.8 Seeley (1993), have assumed a non-linear
–2.0
0 2 4 6 8 10 12 14 16 18 20 22 model and, although both of these models
Temperature (°C) are approximately linear up to 20°C, this
Fig. 10.1. Relationship between air temperature and would encompass the temperatures experi-
chill units used by Hauagge and Cummins (1991) enced by Hamer in Kent. With the fluctuat-
fitted to data from Shaltout and Unrath (1983). ing temperatures during spring, the
relationship between GDH and accumulated
growth can be approximately linear, even if
bud growth rate is not exactly linearly
the magnitude and the duration of the high related to temperature (Cannell, 1989). It is
temperature exposure. The model postu- important to understand that the first visible
lated a dormancy-breaking factor that was external signs of bud development occur
accumulated in buds in a two-stage process, after 30–40% of the GDH accumulation has
the first step being the formation of a pre- taken place (Anderson and Seeley, 1993).
cursor, which was reversible, but, once a Bud temperature is only approximated by
critical portion of the precursor had been screen air temperature. Hamer (1986b) found
accumulated, this was transformed into the that, with light winds and high solar radia-
stable dormancy-breaking factor. This section, buds could be up to 4.5°C above air
ond stage was not reversible. temperature during the day, while
The symptoms of inadequate chilling are Landsberg et al. (1974) showed that bud tem-
seen in delayed and poor bud break, a pro- peratures at night could be 1.5°C below air
longed flowering period, a low proportion of temperature under clear sky conditions. Care
flowering spurs and poor lateral leaf-bud must therefore be exercised in extending
development. Apples are, however, grown detailed studies done by manipulating tissue
successfully in areas that frequently do not temperatures under controlled conditions to
receive adequate winter chilling (e.g. Israel, the orchard environment, where screen air
Brazil). In these areas the trees are treated to temperature may be the only estimate of
enhance bud break. A wide range of chemi- temperature. In some studies, solar radiation
cals have been used for this purpose includ- has been included as an additional term in
ing mineral oil, dinitro-ortho-cresol (DNOC), the relationship between bud development
hydrogen cyanamide, potassium nitrate and and growing degree-days (GDD) to account
6-benzylaminopurine (Lang, 1989). for this effect (Cannell, 1989).
Following the completion of dormancy,
development is dependent on bud tempera-
ture, as altering spring soil temperature by 10.4 Spring Frosts
warming or cooling to give conditions in the
18–26°C or 1–4.5°C range, respectively, com- Although, when fully dormant, apple trees
pared with the control of 4–16°C, did not can withstand 38°C, once dormancy is bro-
influence the time of anthesis of apple flow- ken and active growth has begun, flowers
ers (Hammond and Seeley, 1978). The devel- are very sensitive to low temperatures and
opment of flower buds has been correlated can be killed by late spring frosts. The critical
with the accumulation of growing degree- temperature for damage to the tissues
decreases as the flower buds develop (Table sprinkling, which takes advantage of the
10.6), although some minor hardening is latent heat of freezing. In this case, as ice will
possible if the flowers have been exposed to not form within the tissue until its tempera-
low temperatures and dry conditions prior ture drops below 2°C, the bud is prevented
to the frost event. Within the apple flower, from reaching that temperature by the con-
the styles and ovules are more sensitive to tinual supply of water to the outside of the
damage than the surrounding tissues. bud. As the water freezes, the latent heat of
Following a damaging frost, brown dam- freezing is released to the bud and, assuming
aged tissues can easily be seen by eye after water continues to freeze, the bud does not
sectioning the flower longitudinally with a drop to temperatures that will cause damage.
razor-blade. Cell death results from intracellu- The tree, however, becomes encased in a
lar ice formation causing mechanical damage layer of ice. Although water sprinkling is a
and excessive dehydration (Modlibowska, very effective means of frost protection, it
1975). Less severe damage to the flower can does require an adequate supply of water, as
result in lopsided fruit, due to partial damage the precipitation rates required are up to
to some ovules. Frost rings and russet result- 2.5 mm h1 to provide protection in an air
ing from damage to the epidermal cells, frost of 4°C (Hamer, 1986a). Successful frost
which subsequently produce phellogen and avoidance with water sprinkling therefore
cork formation, are also typical of non-lethal requires both an ample supply of water and
frost events. In some cases the skin splits due free-draining soils.
to ice formation beneath the skin of the The risk of spring frost injury can be fur-
flower receptacle, again resulting in russet ther reduced by the use of late-flowering
development. cultivars or by artificially delaying bloom.
By far the most economical way of avoid- Water sprinkling has been used successfully
ing frost damage is to select sites that, due to to delay blossoming of apples (see Chapter
location, slope and aspect, have a low risk of 20). Bud temperatures can be reduced by
spring frosts, coupled with good cold-air the evaporative cooling of the water. The
drainage (see Chapter 11). There are, how- cooled bud therefore becomes like a wet-
ever, a number of physical steps that are used bulb thermometer, with the greatest cooling
to reduce or eliminate late-spring frost dam- evident in dry environments, such as those
age, all of which rely on the supply of heat to in Utah, where this technique first came to
the sensitive tissues. The flower tissues can prominence. Temperature differences of
be warmed using warm air from heaters (e.g. 20°C were observed between sprinkled and
propane heaters or smudge pots) or by mak- non-sprinkled apple buds in the middle of
ing use of warmer air above the trees (see the afternoon in this dry environment,
Chapter 20). Generally under radiation frost resulting in a bloom delay of 17 days
conditions, there is a temperature inversion – (Anderson and Seeley, 1993). In the more
air temperature near the ground is cooler humid environment of Kent, UK, Hamer
than that several metres above the trees. So- (1986b) found a maximum difference of
called wind machines can be used to stir up 9.2°C between unwetted and wetted buds.
the temperature-inversion layer and raise the In Utah, calculations based on spring tem-
temperature of the flower-buds. These peratures over a 45-year period suggested
machines, however, are limited to regions that a bloom delay of 20 days could have
where there is a strong temperature inver- reduced annual crop losses from 33 to 7%,
sion. An alternative approach is to use water depending on the year. Bloom delay is fre-
Table 10.6. Critical temperatures (°C) for developmental stages of apples.
Green tip Tight cluster Pink Full bloom Post bloom
Temperature for 10% kill 7.8 2.8 2.2 2.2 2.2

Temperature for 90% kill 12.0 6.1 4.1 3.9 3.9
Temperature 227
quently accompanied by a delay in harvest,

although the delay is commonly shorter 50
Pollen-tube growth index

than the delay in blossoming. Consequently,
the technique may increase the risk of cold 40
damage at harvest, and cultivars need to be
30
chosen accordingly in short-season environ-
ments. In terms of timing, water sprinkling
20
is best applied immediately after endodor-
mancy has been completed and stopped at 10
green tip (Anderson and Seeley, 1993). One
of the drawbacks of the technique was high- 0
lighted by Hamer (1981), who found that, 5 6 7 8 9 10 11 12 13 14 15
although water sprinkling delayed blossom- Mean daily temperature (°C)
ing by 14 days in England, the watered buds Fig. 10.2. Effect of daily mean air temperature on
had a higher moisture content and were pollen-tube growth index for a 24-h period. Pollen-
consequently less hardy than unwatered tube growth is completed when the index reaches
trees at the same stage of development. 100. (Figure drawn from data in Williams, 1970.)
In contrast to the situation in regions of
good winter chill, where water sprinkling
may be used to delay flowering, in regions of ‘Ellison’s Orange’ than for ‘Cox’s Orange
inadequate chilling, water sprinkling during Pippin’, following a frost event that resulted
dormancy could enhance the accumulation in equal damage to the flowers.
of chilling hours and, if stopped at the end of In many fruit-growing areas, the set of
endodormancy, could result in earlier flow- flowers is such that the numbers of young
ering of apples. fruitlets have to be reduced quite drastically
to ensure market requirements of fruit size
and quality and to achieve adequate return
10.5 Spring Temperatures bloom the following year. Chemical thin-
ners are frequently used but the efficacy of
Even in the absence of frost injury, tempera- nearly all of these products is temperature-
ture over the blossoming period can have a sensitive (see Chapter 16). Williams and
dramatic effect on fruit set, which in turn Fallahi (1999) suggest that naphthale-
can result in major changes in yield and fruit neacetic acid (NAA) has a temperature opti-
size. Most apple cultivars are self-incompati- mum of 20–25°C, without the risk of
ble, so conditions suitable for pollen transfer overthinning, compared with 15–25°C for
during flowering are necessary. The activity carbaryl and 20–25°C for ethephon, while
of the pollinators, chiefly insects, is Wertheim (2000) reported that the efficacy
enhanced under warm, dry and non-windy of the blossom burner ammonium thiosul-
conditions. Even if pollen is successfully phate (ATS) was much greater at 20°C than
transferred between flowers of compatible at 14°C. Due to the generally higher temper-
cultivars, the rate of pollen tube growth is atures during application of NAA in the
temperature-dependent. Under a mean Pacific Northwest, the rates of chemical
daily temperature of 15°C, pollen tubes take used can be half of those used in the cooler
2 days to reach the ovules (Fig. 10.2), com- eastern states of the USA. The unpre-
pared with 4 days at 13°C and 8 days at 9°C dictability of thinning efficacy still remains
(Williams, 1970). one of the major problems of chemical thin-
Some cultivars (e.g. ‘Cox’s Orange ning, and Wertheim (2000) has recently sug-
Pippin’) are inherently weak in female fertil- gested that more work is needed to
ity and are thus particularly prone to elucidate the effect of weather factors,
adverse weather conditions at flowering including temperature, and their interac-
(Wilson and Williams, 1970). Modlibowska tions on the action of chemical flower and
(1975) quotes a threefold higher yield for fruit thinners.
10.6 Length of Growing Season to 153 days in Kent, UK, over a 22-year
period. Although there was a general trend
The length of the frost-free period effectively of an earlier bloom date resulting in a longer
determines the length of the growing season period of growth, there was also an addi-
for apples in higher latitudes, as the first air tional effect of warm summer temperatures
frosts in the autumn lead to rapid defoliation leading to a shortening of the developmental
of the trees; consequently there is a general period. Each 1°C rise in mean temperature
trend of longer growing seasons at lower lati- during June–August resulted in the harvest
tudes. Taking the data of Kronenberg (1988, date being brought forward by 3.5 days.
1989), a data set that does not include sites
with a major continental influence, there is a
reasonable effect of earlier flowering at lower 10.7 Assimilation and Respiration
latitudes in northern Europe (Fig. 10.3), with
there being, for each degree lower latitude, an The temperature responses of leaf photosyn-
earlier blooming of 2.6 days. This is remark- thesis and respiration are very different. The
ably similar to the 2.5 days calculated by leaf assimilation rate shows a parabolic
Wagenmakers (1991) with a different data set. response to temperature, with a peak at
There is, however, a wide range of cultivar about 30°C, but with a broad shoulder in the
requirements of days from flowering to pick- 15–35°C range (Lakso, 1994). The leaf assimi-
ing. This can be as short as 90 days with lation rate, however, drops off rapidly above
‘Beauty of Bath’ to 200 days for ‘Granny 35°C. In contrast, leaf dark respiration shows
Smith’. Consequently sites with long growing an exponential increase in temperature, so
seasons have a wide range of cultivars avail- that over the 10–30°C range, for each 10°C
able to them, whereas sites with short seasons rise in temperature, the respiration rate
are very limited in the cultivar choice (e.g. in increases by a factor of 2.5 (Lakso, 1994). The
Norway at 60°N the major cultivars are respiration rates of other tissues – wood,
‘Aroma’, ‘Gravenstein’ and ‘Summerred’). fruit and roots – also show an exponential
The length of the period from flowering increase with temperature, although the
to harvest for one cultivar does show some coefficients are somewhat different and vary
variation from year to year. Luton and with the time of year. The respiration rate is
Hamer (1983) found that this period for the sum of two parts, maintenance and
‘Cox’s Orange Pippin’ could vary from 127 growth respiration, where the former is asso-
ciated with the energy required for protein
turnover and the maintenance of ion gradi-
55 ents and the latter is the energy required for
y = −92.8 + 2.58x
new tissue synthesis, which is essential for
Days to full bloom from 1 April
50
growth. It is maintenance respiration that is
45
temperature-sensitive. Consequently, in
40 warmer climates the cost of maintaining cell
35 function increases. Lakso (1994) has drawn
30 attention to the high yields obtained in New
Zealand and explained these partly by the
25
combination of high solar radiation, ensur-
20 ‘Cox's Orange Pippin’ ing high rates of photosynthesis, coupled
‘Golden Delicious’
15 with relatively cool temperatures, ensuring
low maintenance respiration. As an example
10
44 46 48 50 52 54 56 of the effect of temperature on whole-canopy
Latitude (°N) gas exchange, which is the balance between
Fig. 10.3. Relationship between latitude and time respiration and assimilation on the above-
of bloom for ‘Cox’s Orange Pippin’ and ‘Golden ground portion of the tree, Francesconi et al.
Delicious’ derived from data in Kronenberg (1988, (1997) found that CO2 exchange of 4-year-old
1989). potted ‘Royal Gala’ trees decreased by 44%
Temperature 229
when air temperature was increased from early-season apple fruit growth that directly
18°C to 34°C, while it increased by 52% affect fruit size at harvest (Plate 10.1). Fruit
when the temperature was reduced from expansion rates were highly temperature-
18°C to 13°C. The authors acknowledge that responsive only during the cell-division
there was a confounding of air temperature fruit-growth phase lasting c. 40 days after
with vapour-pressure deficit but this does pollination (DAP). Fruit expansion was
frequently occur in the field. Nevertheless, approximately 10 times greater at a mean
the results emphasize the very different temperature of 20°C than at a mean of 6°C.
response of the whole tree to temperature The fruit expansion rate responded to
compared with that of the individual leaf. changes in mean temperature rather than to
the maximum/minimum differential. The
duration of fruit cell division in fact
10.8 Fruit Growth appeared to be inversely related to mean
temperature, being prolonged under cooler
The seasonal pattern of apple fruit growth is conditions. A comparison of mean expansion
defined by an initial 35–50-day period of rates of four cultivars across a gradient from
exponential growth following fertilization, 6 to 20°C also suggested genetic differences.
coinciding with rapidly increasing fruit cell Field studies in three regions of New
number, followed by a more or less linear Zealand over three seasons monitored fruit
growth phase until harvest maturity, during growth of ‘Royal Gala’ under conditions that
which fruit size increases predominantly minimized crop competition. Regression
through cortical cell expansion. An examina- analysis using fruit growth intervals after
tion of the influence of temperature on fruit pollination explored how well temperature
growth and ripening among ten apple culti- accumulation (GDD, base temperature 10°C
vars in four European locations led (GDD10)) was related to seasonal differences
Kronenberg (1988) to propose three sensitive in fruit weight (Stanley et al., 2000). The
periods. Fruit development in the first strongest relationship occurred between fruit
month following flowering and in the weight at 50 DAP and GDD10 at 50 DAP,
period immediately preceding harvest accounting for 71% of the variance. These
responded positively to high temperatures. field-measured fruit growth responses
He concluded that the extended growth closely agree with the controlled-environ-
period between these two stages (probably ment responses from Warrington et al. (1999).
the fruit cell-expansion phase) was insensi- However, temperature accumulation over
tive to temperature. increasing intervals from 30 DAP and includ-
Fruit growth associated with cell division, ing the complete fruit growth period proved
immediately following bloom, appears to be too inaccurate for robust prediction of fruit
very temperature-responsive and important weight at harvest. Early-season temperature
in establishing seasonal fruit-size potential. In effects on fruit growth are clearly integrated
a comparison of fruit growth of four cultivars with other influences, such as competition
over two seasons, larger fruit size at 42 days among fruits, but the relative contribution of
after full bloom (DAFB) was associated with each factor to fruit size at harvest remains
higher temperatures and an increased rate of unquantified. However, it seems likely that,
cortical cell division (Bergh, 1990). However, in commercial production, a reduction in
higher temperatures also enhance fruit set, crop density during the cell-division growth
resulting in a higher fruit number per trunk phase should invoke greatest fruit size
cross-sectional area, which can confound the responses when higher temperatures coin-
exploration of relationships between fruit cide with this early crop thinning.
size at 42 DAFB and size at harvest. There is good evidence that tempera-
In controlled-environment studies, using tures experienced during early fruit growth
methods that avoided crop competition affect the rate and timing of fruit ripening.
effects, Warrington et al. (1999) elucidated There may therefore be an impact on fruit
the fundamental temperature responses of growth and size at maturity if the duration
from bloom to harvest varies greatly. to dissipate heat as well as leaves.

Several controlled-environment studies Consequently the skin temperature of an
showed that fruit-maturity indicators, such apple exposed to strong sunlight can be as
as changes in starch content, firmness, back- much as 13–14°C higher than air tempera-
ground colour, red blush and ethylene pro- ture (Thorpe, 1974). Sunburn of apple fruit
duction, were accelerated by higher was observed by Bergh et al. (1980) when the
temperatures in the first 6 weeks of fruit skin temperature of apple fruit exceeded
growth (Tromp, 1997; Warrington et al., 50°C. This occurred when air temperature
1999). However, each maturity variable did exceeded 36°C. It is not known if tissue sen-
not respond to the same degree. The field sitivity to damage alters with fruit develop-
studies by Stanley et al. (2000) also found a ment or even why some cultivars show less
strong correlation between GDD10 to 30 sunburn damage than others (e.g. ‘Royal
DAP and time from pollination to harvest. Gala’ is less sensitive than ‘Braeburn’ or
Tromp’s study showed the early tempera- ‘Granny Smith’), all of which suggests a
ture influence on ripening to be more influ- fruitful area for future research.
ential than high temperatures applied High summer temperatures can also
around the time of onset of maturity. All reduce the production of flower buds.
these studies were carried out on trees with Controlled-environment studies by Jonkers
their cropping level minimized. (1984) and Tromp (1976) showed that high
Fruit growth and size at harvest appear to daytime temperatures (25–27°C) reduced
be largely unaffected by temperature in the flower formation. In the light of the tempera-
period beyond 40 DAP (Warrington et al., ture effect on net canopy CO2 exchange, this
1999), at least when crop load is minimized. could be interpreted as a lack of available
However, unpublished controlled-environ- carbohydrate, particularly considering that
ment studies have raised the possibility that these trees were grown in controlled envi-
temperature crop-load interactions during ronments, where irradiance levels are often
the fruit-expansion growth phase may be below those experienced in the field.
important in determining fruit development Although the carbohydrate requirement for
and fruit characteristics at maturity (D.S. flower-bud development may be small com-
Tustin, unpublished). pared with that of other sinks, such as fruit,
the tree may allocate carbohydrate preferen-
tially to the other sinks. Shading trees also
10.9 High Summer Temperatures reduces flower-bud production, possibly
through the same mechanism.
Excessively high temperatures can result in
sunburn on the skin of the apple fruit;
although in minor cases this results in a 10.10 Root Growth
clearing of the colour from the epidermal tis-
sues (sun tinting), in more serious cases tis- Rogers (1939) suggested that the onset of
sues can collapse into a brown sunken patch root growth of apple occurs at 6.2°C. The
on the skin, rendering the fruit unsaleable periodicity of root growth, other than during
and more likely to suffer further damage due the winter, seems to be much more related to
to splitting or fungal attack. Current meth- the activity of the aerial part of the tree
ods of control of sunburn rely on reducing rather than to soil temperature. Work in the
the temperature of the fruit skin: (i) by shad- root observation laboratories at East Malling
ing, either by allowing a light leaf cover exte- found a bimodal pattern of root growth, with
rior to the fruit, the use of hail netting or the first flush of growth in May/June fol-
more recently sprays of reflective kaolin; or lowed by a later flush in August/October,
(ii) by sprinkling water on to the apple trees separated by a major peak in shoot growth
and dissipating heat by evaporative cooling. (Atkinson, 1980). There has, however, been
Apple fruit have a considerable thermal considerable work on the effects of high soil
mass and by virtue of their size are not able temperatures on root growth in subtropical
Temperature 231
areas. Gur et al. (1972) reported that root increasing yields. They found that the yield
growth of apple trees was reduced at root of ‘Cox’s Orange Pippin’ was: (i) negatively
temperatures of 30°C and above and serious related to high temperatures in February,
damage to the leaves occurred at root tem- March and April; and (ii) positively related
peratures of 35°C and above, due to the to high temperatures after bloom and again
transport of products of anaerobic respira- in June. Typically in Kent, full bloom of
tion from the roots. ‘Cox’s Orange Pippin’ would be early May.
About 80% of the year-to-year variation in
UK average yields could be explained by
10.11 Shoot and Leaf Growth their relationship, which considering the use
of only one weather data set – that recorded
The effect of temperature on shoot and leaf at East Malling – was particularly encourag-
growth has received much less attention ing. It is important to note, however, that the
than its effects on fruit development. In a majority of the ‘Cox’ crop in England is
study examining the effects of temperature grown in Kent. The negative effect of warm
on shoot development, Tromp (1992) found temperatures before bloom was not associ-
that the production of sylleptic shoots ated with earlier flowering resulting in more
(‘feathering’) was improved as soil tempera- frost damage. The positive effect of high
ture increased to 28°C while air temperature temperatures directly after bloom was
was maintained at 20°C. related to enhanced pollen-tube growth. This
Johnson and Lakso (1985) found that correlation of yield and temperature was fur-
shoot growth of apple was linearly related to ther refined by Miller (1988), also at East
accumulated day-degrees above 4°C, with a Malling (Equation 1), using a data set up to
slope of 0.08 cm GDD1. This rate was quite 1987, where she replaced the mean maxi-
constant among different cultivars, root- mum temperature in February–April by
stocks, tree age and pruning severity. Leaf mean day-degrees above 4.5°C and used a
area per shoot was also linearly related to curvilinear relationship for the long-term
accumulated GDD. In contrast to the results trend of yield:
with shoots, there was considerable variation
Y = 5.27 + 0.5t – 0.009t2 – 2.35DD –
among cultivars in the slope and intercept of
0.51PT + 0.93J (1)
this relationship.
where Y = mean UK yield of ‘Cox’s Orange
Pippin’ (t ha1); t = number of years from
10.12 Synthesis of Temperature Effects 1948; DD = mean day-degrees above 4.5°C
in Models during February, March and April; PT =
number of days to complete pollen-tube
There are two major techniques that have growth; and J = mean maximum tempera-
been used to examine the effect of temperature in June.
ture on tree growth, development and yield: Miller (1988) also examined the causes of
the first uses correlation techniques with the negative effects of warm spring tempera-
long-term data sets of phenology, yield and tures by using mobile greenhouses. Warm
weather data, while the second uses spring temperatures resulted in earlier flow-
controlled-environment facilities specifically ering, as expected, but the effective pollina-
to examine temperature effects directly. Most tion period was also reduced due to poorer
work with apples has made use of the for- ovule longevity, and this resulted in poorer
mer approach, as the size of cropping trees fruit set. Although this work was done with
has limited controlled-environment room ‘Cox’s Orange Pippin’ in England, Lakso
studies to only a few locations (e.g. The (1994) found that the relationship used by
Netherlands and New Zealand). Jackson and Jackson and Hamer (1980) gave a good rela-
Hamer (1980) correlated the average UK tionship to apple yields in New York State,
yield of ‘Cox’s Orange Pippin’ to tempera- suggesting that these effects are more widely
ture, after removing a long-term trend of applicable to other cultivars and locations.
Rather than relying on correlation stud- Temperature during the 4–6 weeks imme-
ies, several more mechanistic models of diately preceding harvest can influence the
apple tree growth have been developed. quality of the fruit at harvest and its storage
The great advantage of such models is that potential. Cooler temperatures result in less
they mathematically relate the growth of all water-core development and a reduced sus-
the parts of the tree to the environmental ceptibility to superficial scald (Bramlage,
variables. Consequently, within the model, 1993) but a higher risk of core flush in ‘Cox’s
temperature and solar radiation can be Orange Pippin’ and ‘McIntosh’ and low-tem-
altered independently at different stages of perature injury in ‘Bramley’s Seedling’,
growth and the outcome predicted. In ‘Cox’s Orange Pippin’, ‘Jonathan’ and
recent work, Lakso et al. (2001) used a ‘Yellow Newtown’ (Sharples, 1984). Less
model to predict net CO2 assimilation by scald after storage was observed where fruit
apple trees in New Zealand, Washington had received 100–150 h below 10°C before
State and New York State. The authors harvest and little scald developed when the
draw attention to the mild temperatures in fruit had received more than 150 h (Merritt et
New Zealand extending the leaf-area dura- al., 1961). In the UK, however, increased inci-
tion after the harvest period. In some of his dence of scald has been related to increasing
earlier correlation studies between temper- soil-moisture deficit from late July to early
ature and annual yields in New York, September with the cultivar ‘Edward VII’
Lakso (1994) found that mean temperature (Sharples, 1984). Sharples adds a key general
between harvest and leaf fall had a positive comment that the incidence of storage disor-
effect on yield in the following season, pos- ders is frequently related to fruit maturity at
sibly via extended leaf-area duration. harvest, and relationships to temperature,
Wagenmakers (1996) has modified the crop for example, may be confounded with effects
growth model SUCROS, developed by the on fruit maturity if picking date is not
group at Wageningen, for apple trees. The adjusted accordingly.
results from this model predicted that
warm springs were favourable for a rapid
development of leaf area, resulting in 10.14 Conclusions
increased light interception and CO2 assim-
ilation. In contrast, higher temperatures in Temperature has a profound effect on all
mid- to late summer before harvest aspects of apple tree physiology. At the
resulted in high maintenance respiration extremes, high or low temperatures can
and reduced fruit growth. cause death, while, at intermediate tempera-
tures, all physiological processes are affected.
It must not be forgotten, however, that tissue
10.13 Fruit Quality temperatures within the plant vary widely
from exposed to shaded tissues and from tis-
Although fruit colour development on red sues above ground to tissues underground,
cultivars of apples is strongly dependent on where there is also a range of soil tempera-
cultivar, fruit maturity, light, nutrition and tures. The response of the whole plant to
crop load, there is also a pronounced effect of temperature is the result of the effect of tem-
temperature (Saure, 1990). In general, cooler perature on each part of the plant and the
night temperatures favour the development interactions among those tissues. With the
of red colour – for example, Creasy (1968) changes in climate associated with global
found an inverse relationship between aver- warming, it is even more imperative to eluci-
age temperature and anthocyanin formation date the response of the apple tree to temper-
in ‘McIntosh’ apples. The relationship ature to be able to predict the effect of
between colour formation and temperature increasing temperatures in order to guide
also seems to vary between experiments orchardists in their choice of cultivars and to
done with detached fruit and those done alert them to a likely increased or decreased
with whole trees with attached fruit. incidence of problems.
Temperature 233
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11 Selecting the Orchard Site, Site

Preparation and Orchard Planning and
Establishment
John A. Barden1 and Gerry H. Neilsen2

1Department of Horticulture, Virginia Polytechnic Institute and State University,
Blacksburg, Virginia, USA; 2Agriculture and Agri-Food Canada, Pacific Agri-Food
Research Centre, Summerland, British Columbia, Canada
11.1 Selecting the Orchard Site 238

11.1.1 Geographical and climatic considerations 238
11.1.2 Site selection: temperature 239
11.1.3 Winter injury 241
11.2 Soil Considerations 242
11.2.1 Soil physical properties 242
11.2.2 Soil topography 246
11.2.3 Soil chemical properties 246
11.2.4 Soil chemical contamination 248
11.3 Site Preparation 248
11.3.1 Tillage 248
11.3.2 Landscape modification 249
11.3.3 pH adjustment 249
11.3.4 Other soil amendments 252
11.4 Replant-site Preparation 253
11.4.1 Fumigation 253
11.4.2 Alternatives to fumigation 254
11.5 Orchard Planning 255
11.5.1 Cultivar selection 255
11.5.2 Rootstock selection 255
11.5.3 Row orientation 258
11.6 Tree Establishment 258
11.6.1 Tree quality 258
11.6.2 Tree planting 259
11.6.3 Tree support 260
11.6.4 Pruning at planting 261
11.7 Summary and Conclusions 262

238 J.A. Barden and G.H. Neilsen
11.1 Selecting the Orchard Site and Georgia. In Mexico, commercial quanti-
ties of apples are grown at high elevations in
11.1.1 Geographical and climatic the northern states of Chihuahua, Durango
considerations and Coahuila (Herrera, 1984).
The cultivated apple has its origin in the

11.1.1.2 Proximity to oceans or lakes
Republic of Kazakhstan (see also Chapter 1),
from which it has spread throughout the Another factor of considerable importance is
world (Hokanson et al., 1997). It is classified proximity to large bodies of water, particu-
as a temperate fruit tree, which indicates that larly when the prevailing winds come over
it is deciduous and requires an extended cold the water. Apple industries exist in Ontario,
period (chilling requirement) for buds to Canada, primarily in the southern regions of
break for both foliation and flowering. Apple the province along Lake Erie and Lake
trees can survive in many parts of the world, Ontario. Likewise, apple areas exist on the
but most are grown between latitudes 30° leeward sides of the Great Lakes in Michigan
and 50° north and south. The primary limit- and New York. Nova Scotia has a significant
ing factor in apple production at latitudes apple industry because of the moderating
lower than 30°, north or south, is the lack of effects of the Bay of Fundy and the Atlantic
adequate chilling in winter to break rest Ocean. There is very little apple production
(endodormancy). Additionally, the exces- in the middle Canadian provinces of Alberta,
sively warm temperatures at such latitudes Saskatchewan and Manitoba because the cli-
have negative effects on fruit colour and mate is continental; their winters are so cold
quality. At greater than 50° latitude, north or that they severely restrict apple production.
south, the major limitations related to tem- There is an important apple industry in inte-
perature are damage by the coldest nights of rior valleys of southern British Columbia
winter and inadequate length of growing sea- because of the moderating influence of the
son. At all latitudes, there are sizeable effects Pacific Ocean and large valley lakes, such as
of other factors, such as elevation and prox- the Okanagan (Plate 11.1).
imity to large bodies of water. Bodies of water serve two vitally impor-
tant roles in minimizing frost damage. In the
spring, water warms up more slowly than
11.1.1.1 Elevation
the adjacent land surfaces. The prevailing
Temperatures are dramatically affected by winds are cooled as they pass over the cold
elevation; a rule of thumb is that day and water in the spring. As the cooler air passes
night temperatures decline by about 3°C for over the adjacent apple trees, bud develop-
each 500 m increase in elevation. A striking ment is delayed, thus decreasing the proba-
example of this effect on temperatures is bility of spring frost injury. In the autumn,
found in South America. Belem, Brazil, is 19 the water cools more slowly than the land
km from the equator at an elevation of about and thus air masses are warmed as they
10 m and has a mean annual temperature of pass over it. This warmer air delays the
29°C. Quito, Equador, also about 19 km from onset of early-autumn freezes. This lake
the equator but at an elevation of 2835 m has effect is of consequence for only a few kilo-
a mean annual temperature of 13°C. The metres from the lake but accounts for the
effect of elevation is present regardless of lat- large concentration of orchards and vine-
itude and plays a vital role in determining yards along the leeward sides of lakes or
what crops can be grown successfully. other large bodies of water. Since relatively
Therefore, as latitude decreases, commercial mild autumn frosts (1 to 3°C) are not
apple orchards are planted at higher eleva- particularly detrimental to leaves and
tions. For example, in the eastern USA, apple apples remaining on the tree, this extension
orchards south of Virginia are mostly in the of the growing season is much more valu-
mountains, such as in western North able to a grape grower, whose crop is
Carolina and north-western South Carolina severely damaged by even a light frost.
Selecting the Orchard Site 239
11.1.1.3 Chilling requirement 11.1.1.4 Length of growing season

The apple is a fruit tree in the general cate- The length of the growing season, also
gory of temperate plants, which are charac- referred to as the number of consecutive
terized as requiring an annual cold period to frost-free days, varies greatly with latitude
satisfy their ‘chilling requirement’. If the and depends upon a number of factors.
chilling requirement is not satisfied, the buds There is a considerable impact of large bod-
will not open; if the chilling requirement is ies of water on spring, autumn and winter
partially met, the buds will open sporadi- temperatures. In general, lakes extend the
cally and both the bloom and harvest peri- growing season within a few kilometres of
ods will be abnormally extended. Most apple the leeward side. Another very important
cultivars require between 1000 and 1200 h of factor is elevation, in this case relative to
chilling in the range of 4–7°C. Temperatures sea level rather than above the adjacent ter-
below 0°C or above about 10°C do not pro- rain (as will be described below under
vide chilling and may actually negate previ- spring frost potential). In very poleward
ously accumulated chilling hours. Because of countries, such as Denmark, Norway and
this, the chilling requirement of a particular Sweden, the moderating effect of the sur-
cultivar may well be satisfied earlier in a rounding oceans offers some protection
middle latitude than in a more poleward lati- from extreme winter cold, but the growing
tude. In some parts of the world where win- seasons are very short. The number of days
ter temperatures are so warm as to barely from blossoming to harvest for apple culti-
meet the chilling requirement (as in parts of vars ranges from 75 or 80 to over 200.
California), the presence of frequent morning Short-season cultivars can be grown
fog delays the daily rise in temperature of widely, but those requiring more than 180
buds, thus enabling the accumulation of ade-
days are restricted to regions with long
quate chilling hours.
growing seasons.
In warm areas where the chilling require-
ment is not met, rather extreme treatments
can induce the buds to grow despite the lack
of chilling (Edwards, 1987). According to 11.1.2 Site selection: temperature
Janick (1974), cultivars including ‘Rome
Beauty’ and ‘McIntosh’ as well as local culti- In the selection of a site on which to estab-
vars can be grown in areas of Indonesia (east lish an orchard, climatic considerations are
Java) at latitudes of less than 10°S and eleva- of utmost importance. In the remainder of
tions of 700–1200 m. The maximum, mini- this chapter we discuss climate from a some-
mum and mean annual temperatures are 31, what local perspective. Arguably, the most
22 and 26°C, respectively. Through the use of critical climatic factor to consider in select-
complete defoliation approximately 1 month ing an orchard site is temperature and,
after harvest, a new growth and fruiting among the several aspects of temperature,
cycle is initiated. Flowering occurs approxi- none approaches the importance of the
mately 1 month after the trees are defoliated. likelihood of spring frost damage.
The cycle is repeated to induce two crops per Unfortunately, spring frost damage to apple
year. In subtropical regions, the chilling flowers and young fruit is an all-too-fre-
received is inadequate and various treat- quent occurrence (Plate 11.2). There have
ments have been developed to supplement been a great many approaches devised to
the chilling received. In addition to the selec- minimize spring frost damage (see Chapter
tion of low-chilling cultivars, such as ‘Anna’ 20), but, by an immense margin, the best
and ‘Ein Shemer’, several chemicals have frost-control technique is good site selection!
proved to be at least somewhat effective. There are obviously many other considera-
These include mineral oil, dinitro-o-cresol, tions, such as soil, in selecting a good
potassium nitrate, thiourea and cyanamide, orchard site, but, if the site is ‘frosty’, none
as well as gibberellins and cytokinins (Erez, of the other attributes of the site will over-
1987). come this one all-important weakness.
11.1.2.1 Radiational freezes normal situation during the day when the air
temperature declines with increasing alti-
Considerable information can be gained by
tude. The ‘top of the inversion’ is the altitude
visually evaluating the site under considera-
at which the air is warmest. Both wind
tion. Most important is its elevation, not machines and helicopters are used to mix the
above sea level but above the adjacent ter- warm air in the upper portion of the inver-
rain. The vast majority of frost damage sion with colder air at tree level in the
occurs under radiational freeze conditions. orchard (see also Chapter 20).
On relatively clear, calm, spring nights, heat On a commercial scale today, orchard
is lost by radiation from plants, soil and other sites are selected on the basis of minimal
surfaces exposed to the cold sky. The lower danger of radiational frost damage, but they
layers of air are cooled by losing heat, by con- must also be on a slope that does not pre-
duction, to these exposed surfaces, which clude the use of modern equipment. Since
have dropped below air temperature. As air apple trees thrive in a permanent-sod soil-
is cooled, it becomes more dense (heavier) management programme (often supple-
and therefore settles on the ground, flowing mented with a herbicide strip), such
ultimately into low areas; this settling of cold orchards can be successfully grown on
air has led to the term ‘frost pocket’, used for steeper slopes than peaches, which are usu-
low-lying areas that are particularly subject ally grown with at least some cultivation.
to frost damage (Fig. 11.1). The degree of slope that can be tolerated will
vary with row spacing, tree size and espe-
cially the type of equipment available (Plate
11.1.2.2 Temperature inversions
11.3). In different parts of the world, these
As lower air layers lose heat to cold surfaces, climatically ideal fruit sites may have some-
these layers become colder than the air what different characteristics. In many
above, so air temperature increases with alti- regions, orchard sites are selected on sloping
tude. This phenomenon is called a ‘tempera- land above large valleys, on knolls in rolling
ture inversion’ because it is the reverse of the terrain and on terraces above rivers.
4 4
3 3
Temperature (°C)
2 2
1 1
0 0
–1 –1
–2 –2
–3 –3
Cold air Frost pocket
Fig. 11.1. During a radiational freeze, layers of cold air along the ground flow downhill, settling into low-
lying areas called ‘frost pockets’.
11.1.2.3 Advective freezes length and declining temperatures. Sub-

freezing temperatures are required to effect
Apple crops are sometimes lost to cold tem-
the later stages of the hardening process.
peratures associated with an ‘advective
Another factor that contributes to the occur-
freeze’, which is a cold event resulting from
rence of winter injury is the variation in tem-
the invasion of a large cold air mass accompa-
peratures before the cold event. A freezing
nied by high winds (see also Chapter 20).
event preceded by a warm period is far more
Since such conditions preclude the formation
damaging than continuously cold tempera-
of a temperature inversion, wind machines,
tures. The development of hardiness is a grad-
helicopters and other methods dependent
ual process that proceeds over a period of
upon an inversion are ineffective. Although weeks, but even a temporary warm period late
much less frequent than radiational freezes, in winter can lead to rapid loss of hardiness.
the occurrence of an advective freeze is devas- For this reason, winter injury to apple trees is
tating because there is little that can be done more common in winters with wide tempera-
to lessen its impact. Furthermore, the sites ture fluctuations than in winters with compa-
selected to minimize danger from radiational rably low but continuously cold temperatures.
freezes are particularly vulnerable to advec-
tive freezes because of increased exposure.
11.1.3.2 Trunk injury
Whether it is a young tree damaged in the
11.1.3 Winter injury autumn or an old tree damaged in midwin-
ter, the injury is most often to the trunk and
Winter injury to apple trees can vary both in lower scaffold limbs. These are the last tis-
when it occurs and in what trees are injured, sues to harden and therefore the most likely
depending on a variety of factors (see Chapter to be injured. Injury is frequently apparent as
10). One type of winter injury typically occurs a splitting of the bark (Fig. 11.2). If the bark is
in mid- to late winter to mature bearing trees reattached to the trunk or branch soon
that have reached their maximum hardiness, enough to avoid death of the vascular cam-
but are injured because of the extreme cold bium by desiccation, the tree can be saved. If,
(often in the range of 30° to 35°C). With however, the cambium dries out on a large
bearing trees, previous stress resulting from a part of the trunk, the only way to save the
particularly heavy crop, drought, nutrient tree may be to utilize either bridge grafting or
deficiency or premature defoliation can lessen inarching (Hartmann et al., 1990). With bridge
midwinter cold-hardiness. A second type of grafting, the goal is to replace the trunk; with
injury consists of damage in late autumn to inarching, the goal is to replace both the
young trees that have not yet reached their trunk and the root system. In severely cold
maximum hardiness. Because of either heavy periods, the entire trunk may split open,
or late nitrogen applications, young trees tend which allows little hope of survival.
to grow relatively late in the summer and are
therefore delayed in reaching maximum har-
11.1.3.3 Other factors
diness compared with an older bearing tree,
which ceases vegetative growth earlier in the There is considerable variation in the cold-
summer. hardiness of both rootstocks and scion culti-
vars, and this can have important practical
implications. The rootstocks selected in
11.1.3.1 Hardening
breeding programmes such as those in
The physiological processes involved as Poland (P series) and Russia (Budagovsky
perennial plants develop cold-hardiness series) have been selected to a large degree
have been studied extensively, but the exact on the basis of cold-hardiness because win-
mechanisms remain elusive. The hardening ter survival is such a vitally important issue
process starts only after vegetative growth in those areas (Ferree and Carlson, 1987).
has ceased and is induced by decreasing day Winter survival is uppermost in the selec-
11.1.3.4 Root injury

The roots of apple trees are considerably less
cold-hardy than are the above-ground por-
tions. Much of this difference is because roots
do not go into rest and are active during win-
ter when soil temperatures are above a mini-
mum of about 6–7°C. Secondly, since roots are
not exposed to temperatures as low as are the
above-ground portions, they do not develop
nearly as much hardiness. If an apple root is
exposed to the same gradually declining tem-
peratures as the above-ground parts of the
tree, the hardiness of the root can become
equal to that of a branch. A continuous snow
cover provides considerable protection for the
roots of apple trees by insulating the soil from
excessive heat loss. Because snow is made up
of about 80% entrapped air, it is an excellent
insulator. Winter injury to the roots of apple
trees is far more likely to occur in areas with
an ‘open winter’ without snow cover than in
more northern regions, which may be colder
but have more snow on the ground.
11.2 Soil Considerations
General observations from many fruit-grow-

ing regions indicate that apple trees grow
well on a range of soils. However, the extra
effort and cost required to manage difficult
soils indicate that considerable effort should
be devoted to investigating and understand-
Fig. 11.2. The splitting of bark on the trunk of apple
ing limiting soil properties and their poten-
trees is a common symptom of winter-cold injury. It
may occur from a moderately cold temperature tial amelioration at a site prior to incurring
(10°C) in mid-autumn, prior to the development the additional costs associated with planting.
of maximum hardiness, or from severe cold (30°C)
in midwinter.
11.2.1 Soil physical properties
tion criteria of apple breeders in areas with It is generally acknowledged that poor soil
very cold winters, such as Minnesota; for physical properties require paramount con-
apple breeders in warmer regions, winter- sideration because they are more difficult
hardiness is of much less concern. and sometimes impossible to improve as
Severity of winter injury within a region compared with inadequate soil fertility and
tends to vary little due to local factors such chemical conditions.
as elevation, because winter cold events are
usually associated with windy conditions,
11.2.1.1 Soil texture
much like an advective freeze in the spring.
In winter there are therefore not the large The texture of a soil refers to the proportion
microclimatic effects that are so important of different mineral particle sizes in the soil.
during radiational freezes in the spring. Size classifications from smallest to largest
include clay (< 0.002 mm), silt (0.002– pores and fissures throughout the soil pro-
0.05 mm), sand (0.05–2 mm) and gravel file, allowing easier air and water movement
(2–80 mm). The percentage by weight of sand, (Fig. 11.4). A stable structure with a wide
silt and clay in a given soil is often represented range of pore sizes is desirable for apple pro-
in a texture triangle from which soils can be duction. Serious growth problems can arise
grouped into similar textural classes ranging when compacted layers with high bulk den-
from fine to coarse (Fig. 11.3). The textural sity and associated reductions in air-filled
grouping influences several soil properties voids occur in the main rooting zone. Such
important to apple production, including layers can arise near the surface in traffic
potential water-storage capacity, perviousness rows of old orchards subjected to frequent
to internal water transmission, aeration and passes of tractors and sprayers under moist
nutrient-exchange capacity (Table 11.1). A spe- soil conditions. Some soils can also contain
cial consideration regarding both soil texture naturally compacted subsoil ‘hardpans’
and soil structure is the ability of the soil at a resulting from layers enriched in calcium,
site to support the year-round passage of iron or clay particle sizes.
orchard equipment (‘trafficability’). Soils
should be able to withstand compaction and
11.2.1.3 Soil depth
be free of surface flooding. Large gravel, cob-
ble and stones (> 2 mm diameter) can improve Apple tree performance is affected by the
drainage and warming of some fine-textured depth of the soil to which unrestricted root
soils but can be serious impediments to soil growth can occur. Possible barriers to root
cultivation and even tree planting. development include seasonal or year-round
high water-tables, compacted layers, bedrock
or abrupt textural changes to much finer or
11.2.1.2 Soil structure
coarser particle size in the subsoil. The depth
In most soils, individual particles of sand, of unrestricted root penetration by apple can
silt and clay are aggregated into larger-scale be quite variable, with a 1–2 m depth most
structural units, which create a network of common, although most roots are generally
located within the surface 0.8 m depth
(Atkinson, 1980). A greater potential rooting
Fine depth can enhance tree growth and produc-
tion by increasing tree access to nutrients
100 Moderately and water. This can be particularly impor-
fine
90 tant in coarse-textured soils, which often
Medium have limited nutrient- and water-holding
80
capacities (Table 11.1). Although some of the
70 Moderately limitations of shallow soil depth can be over-
coarse
60 come by effective irrigation or soil amend-
Coarse ment, it is a well-known phenomenon that
Clay (%)
50
severe restrictions in rooting volume reduce
40 tree vigour and fruit size in spite of modifi-
cations to improve nutrient and water sup-
30
ply to the plant. It is therefore important to
20 have a knowledge of effective soil depth
10 prior to planting.
Shallow soil depths can be less limiting to
0
0 10 20 30 40 50 60 70 80 90 100 apple tree performance for scions on rootstocks
Sand (%) that are already naturally dwarfing. However,
Fig. 11.3. Percentage of clay and sand in the main care must be taken to optimize soil physical
textural classes of soils; the remainder of each class and chemical conditions in what is a more lim-
is silt (adapted from Association of British Columbia ited soil volume that offers less buffering
Grape Growers, 1984). against various environmental stresses.
Apples - Chap 11
244
21/3/03
Table 11.1. General characteristics of main soil textural groups (adapted from Association of British Columbia Grape Growers, 1984).
2:56 pm
Typical nutrient
Typical water exchange capacity
storage capacity (cmol (positive Trafficability
Textural group Soil textures (cm m1) Perviousness Soil aeration charge) kg1) (when wet)
Page 244
J.A. Barden and G.H. Neilsen
Coarse Gravel (G) Very low–low Rapid (soil remains High Very low Good
Sand (S) (2–10) wet for only hours (< 5)
Loamy sand (LS) after wetting)
Moderately coarse Sandy loam (SL) Moderate Rapid–moderate High–moderate Low Good
(10–14) (5–15)
Medium Loam (L) Moderate–high Moderate (soil remains Medium Medium Moderate
Silt loam (SiL) (17–21) wet for days after (15–25)
Silt (Si) wetting)
Moderately fine Clay loam (CL) High Poor (soil remains wet Medium-low High Poor
Silty clay loam (SiCL) (≥ 20) for weeks after wetting, (20–50)
Sandy clay loam (SCL) many small pores)
Fine Silty clay (SiC) High Poor Low High Very poor
Sandy clay (SC) (≥ 20) (20–50)
Clay (C)
Heavy clay (HC)
Granular
Granular and
blocky Coarse structured
puddled clay – worked
when wet
Platy
Weak blocky Plough pan
Blocky
Very large prismatic

tightly fitting
Prismatic Very slow water
breaking to movement
blocky
Fig. 11.4. Example of well-structured soil (left) and poorly structured soil (right) (adapted from Davies et al.,
1972).
11.2.1.4 Soil water relations space for the aeration of roots. PWP is the
The capacity of soils to store water available water content at which plants can no longer
to plants varies considerably with soil tex- extract adequate water to regain turgidity in
ture (Fig. 11.5). Water availability is usually the dark. Local soil survey reports often con-
expressed as the difference in water content tain available water-content values for vari-
between field capacity (FC) and the point at ous soils by measuring the difference in
which plants wilt permanently (PWP). FC is water content between soils equilibrated at
the water content after drainage of saturated pressures between 0.03 MPa (FC) and
soils has become negligible. The drained 1.5 MPa (PWP). Such information can be
pore space under this condition provides air valuable when selecting planting sites for
40 0.4
Water content (cm cm–1 of soil)
Total water
In situ
30 field 0.3
Volume soil water (%)
Available water
capacity
20 0.2
Permanent
10 wilting percentage 0.1
Unavailable water
0 0
Sandy Sand
Loam Silt Clay Clay
loam loam loam
Fig. 11.5. Relative amounts of available and unavailable water in soils ranging in texture from sands to
clays. Amounts of water expressed as percentages of soil volume (left) and cm of water per cm of soil depth
(right). (From Kramer and Boyer, 1995.)
apple orchards. Growth limitations from As previously indicated, landscape location

inadequate water-storage capacity or the can result in more favourable microclimate
need for supplemental irrigation is more conditions, which can enhance orchard per-
likely to occur on coarse-textured soils. formance by reducing the risk of spring-
Also important for apple trees is good soil frost damage.
aeration for proper root functioning. Aeration
is not often a problem in coarse-textured soils
but can seriously limit growth in fine-tex- 11.2.3 Soil chemical properties
tured soils. Poor internal drainage through
soil profiles can result in excessive waterlog-
11.2.3.1 Nutrient content
ging, depriving roots of oxygen and creating
saturated soil conditions favourable for the The availability of many plant nutrients,
development and spread of pathogenic fungi, including major elements, such as phospho-
such as Phytophthora root and crown rots, rus, potassium, magnesium, calcium and
especially on susceptible rootstocks. sulphur, and trace elements, such as zinc,
Good soil drainage is thus preferable manganese, boron, iron and copper, can be
when locating apple orchards. Sites suscepti- readily determined in various extracts from
ble to waterlogging or even surface ponding representative composite soil samples col-
of water can occur in low-lying areas where lected at a prospective orchard site. Since
water tables are high year-round or only apple is relatively deep-rooted compared
during periods of excess moisture, as in the with many annual crops, composite sam-
spring after snow-melt or when evapotran- ples need to be collected to a sufficient
spiration is low and precipitation high. Such depth to characterize the anticipated major
locations may occur because of restriction of rooting zone. Thus, in addition to samples
water movement through impermeable sub- collected from the surface layers (to 20 cm
soils or may result from seepage of water depth), useful information can be obtained
(and sometimes excessive fertilizers and from samples collected to represent 20–40
other soluble salts) from other locations in and 40–60 cm depths. The usefulness of the
the landscape. Although it is possible to rem- values determined often depends upon the
edy inadequate drainage at some sites by extent of local research to calibrate tree
installation of tile drains, this can be an response to fertility parameters. Usually
expensive solution. much less information is available for
apples than for annual agronomic and hor-
ticultural crops. However, it can be
11.2.2 Soil topography extremely useful to determine if a site has
exceptionally low or high availability of
The location of a potential orchard site important plant nutrients and how avail-
within the general landscape can affect sub- ability of these nutrients changes with
sequent orchard performance. Flat or gen- depth. Ameliorative fertilizer applications
tly sloping sites are ideal for the operation can thus be made prior to planting when
of equipment. Other considerations include relatively immobile nutrients, such as phos-
location with respect to prevailing winds phorus and potassium, can be ploughed
and local air flows. Exposed ridges and into the potential root zone. Phosphorus
narrow valleys subjected to persistent seems to be a particularly important
strong winds can result in disruption in the nutrient for establishing apple trees.
timing and accuracy of spray application Improvement in first-year growth of apple
and can even distort tree shape. Tree wind- trees has been reported following applica-
breaks have been effectively planted as bar- tions of relatively high rates of especially
riers to reduce wind effects on orchards, well-mixed ammonium phosphate fertilizer
although edge rows of apple trees are sus- compounds directly in the planting hole or
ceptible to the competitive effects of vigor- fertigated directly into the zone of newly
ously growing trees in nearby wind-breaks. emerging roots (Neilsen, 1994). Nitrogen,
however, is the major fertilizer to be applied most important soil factor affecting man-
in most orchards but, at present, N-fertilizer ganese solubility, waterlogged soils, regard-
recommendations are adjusted on the basis less of pH, can also have an increased
of plant response or leaf analysis rather incidence of manganese-induced internal
than based on soil N assessment. bark necrosis (Grasmanis and Leeper, 1966).
Soluble aluminium is also highly toxic to
most plants, even at a low concentration, act-
11.2.3.2 pH
ing to inhibit root elongation and branching
An ideal soil pH range (as measured in (Scott and Fisher, 1989). Aluminium is toxic
water) for apple is 6.5–7.0, although there to apple seedlings at concentrations as low
has been limited documentation of this infor- as 6.5 106 M (175 p.p.m.) in solution
mation. It is, however, known that below pH (Kotze et al., 1976).
5.5 the solubility of manganese and alu- In alkaline soils, where pH values exceed
minum ions increases rapidly in most soils. 7.0, other growth limitations have been
Manganese toxicity has been associated with observed. For example, trees planted on cal-
internal bark necrosis (Fig. 11.6) and reduced careous soils, where pH is frequently
vigour of newly planted trees (Hoyt and 7.5–8.5, can have growth-limiting nutrient
Neilsen, 1985). Manganese toxicity problems deficiencies resulting from decreased avail-
are especially serious for ‘Delicious’ and cul- ability of phosphorus and several trace ele-
tivars with ‘Delicious’ parentage but have ments, including iron, zinc and manganese.
been observed on several other cultivars, Alkali soils have pH values exceeding 8 and
including ‘Golden Delicious’ and ‘McIntosh’ high sodium contents, which adversely
(Berg et al., 1958). Although a low pH is the affect soil structure, resulting in dense and
poorly aerated soils.
As will be discussed later, considerable
research has been conducted on ways to
overcome adverse soil pH conditions. By far
the best time to make such adjustment is
prior to planting, when concern for existing
trees will not limit the ability to work
amendments to sufficient depth.
11.2.3.3 Salinity
Saline soils can exhibit a wide range of basic
pH values and can have high concentrations
of salts, such as sodium, chloride and boron,
which can be toxic to apple trees. Soils with
high salinity values in what will be the major
rooting zone of a proposed apple orchard
can limit tree establishment. Considerable
information available from research organi-
zations (such as the USA’s Salinity
Laboratory in Riverside, California (Maas,
1987)) indicate that apple, like most peren-
nial woody fruit crops, is susceptible to
yield and growth reductions as soil salinity
increases. Soil salinity is usually measured
as the electrical conductivity of saturated
Fig. 11.6. ‘Starkrimson Delicious’ apple tree on extracts of composite soil samples represent-
M.26 rootstock exhibiting severe internal bark ing the main root zone. Values are expressed
necrosis resulting from manganese toxicity. as decisiemens per metre (dS m1) and val-
ues much exceeding 1 dS m1 (approxi- 11.3 Site Preparation

mately 640 p.p.m. (mg l1) salt in soil solu-
tion) have been reported to reduce the yield There are a number of actions that can be
and growth of apple. Excessive soil salinity undertaken to improve the suitability of a
can occur in landscape locations in semiarid site for orchard establishment. However, the
regions where seepage waters accumulate, cost of such activities needs to be accounted
but can also be induced in smaller soil for when deciding where to plant.
volumes by over-application of soluble
fertilizers to the roots of newly planted
apple trees. 11.3.1 Tillage
Normally, tillage operations are undertaken

11.2.4 Soil chemical contamination to about a 20 cm depth in orchards prior to
planting in order to produce desirable con-
Accumulation in the soil of persistent chemi- ditions for the growth of the planted trees.
cal residues from previous land-use activities Successful tillage may act to maintain soil
has the potential to adversely affect growth infiltration capacity and aeration, to mix
of newly planted trees. For old orchard sites, pre-plant amendments and fertilizer
for example, high lead arsenate concentra- throughout the main rooting zone and to
tions in the root zone can be phytotoxic to control weeds. Increasingly, less emphasis is
apple trees (Benson, 1976). Lead arsenate placed upon tillage for the purpose of weed
(and related compounds) was widely used control, in part because of possible detri-
as an insecticide, particularly for control of mental effects on soil structure, but also
codling moth (Cydia pomonella L.) for about a because of the effectiveness of weed control
50-year period, ending about the time of the via herbicides.
introduction of the synthetic organic insecti- Sometimes mechanical manipulation of
cide dichlorodiphenyltrichloroethane (DDT) soils 15–25 cm below the normal tillage
following the Second World War (Peryea, depth is warranted. These activities, also
1998). In Washington State, growth problems referred to as deep ploughing, chiselling,
with young apple trees are predicted when subsoiling or ripping, are intended to break
total arsenic concentrations in the root zone up any compacted or impermeable layers
exceed 50 p.p.m. (Benson, 1968). Remedying within the root range of young trees. This
such a situation can be difficult without the should increase infiltration and pore size for
addition of uncontaminated soil, particularly drainage and water storage, allowing
in and near the planting hole where newly improved root growth and leaching of
forming roots emerge. excess water and salts. The effectiveness of
It is also known that applications of the procedure varies with soil moisture con-
residual herbicides, such as terbacil (3-tert- tent at the time of ripping, being more effec-
butyl-5-chlor-6-methyluracil), diuron (3- tively performed on dry soils. It is
(3,4-dichlorophenyl)-1,1-dimethylurea) or physically much easier to plant into soils
simazine (2-chlor-4,6-bis(ethylamino)-s-tri- after deep ripping. However, ripping as a
azine), above recommended rates can general preplant orchard practice is proba-
decrease the subsequent vigour and yield of bly not justified, as limited growth improve-
young apple trees (Hogue and Neilsen, ments may not warrant the added expense.
1988). It is best to be alert to this possibility Even on soils and for other crops where
by knowing the history of previous weed deep ripping is more commonly under-
control at a prospective planting site. taken, increases in growth or yield or
These examples show that it is useful to improvements in soil physical properties
investigate the implications of previous land have been inconsistent, sometimes tempo-
use at a site prior to making the decision to rary and frequently disappointing (Wild,
replant. 1988).
11.3.2 Landscape modification row ground-cover strips, which reduce

water runoff and soil erosion.
11.3.2.1 Levelling
After clearing the land of previous vegeta- 11.3.2.3 Terracing
tion or old orchard trees, it may be useful
Terracing is usually considered an extreme
and relatively inexpensive to level an irregu-
form of land shaping. Although practised for
lar surface. Thus surface drainage depres- thousands of years in many parts of the world
sions may be infilled and a flatter field to conserve moisture and reduce erosion,
established for operation of orchard equip- establishment costs are high. The procedure
ment and irrigation systems. (With the involves the construction of broad benches
advent of pressure-compensating emitters, across the steep slopes (usually 20–30°) of
slope requirements are less exacting for rolling land. Such practices are usually con-
modern low-pressure irrigation systems.) sidered only when other soil-conservation
The degree of levelling depends upon the methods are ineffective. Construction details
original land surface. Large gulleys have are contained in many standard soil- and
been successfully infilled to create larger water-engineering texts (Schwab et al., 1966).
contiguous areas of level land. However, A variation of the complete terracing method
problems can arise, since planted apples fre- is the construction of a single diversion
quently grow poorly on exposed subsoils or embankment up-slope of an orchard in
new fill, which can have poor fertility or humid climates to prevent slope runoff water
structure. Pre-plant land modification can be from flooding orchard land.
expensive but has been successfully applied
prior to planting. For example, the ‘Tatura’
soil management system involves pushing 11.3.2.4 Drainage modification
topsoil into ridges in order to increase soil The benefits of improved soil drainage can
depth and improve drainage within the tree allow successful production of apples on
row when shallow top soils overlie compact soils and in locations where waterlogging
subsoils (Baxter, 1977). Similarly, in south- and poor aeration would normally limit the
eastern USA, subsoils have been reshaped development of orchards. For example,
and levelled with removed topsoil being much apple production in The Netherlands
respread to create a new soil surface with has proceeded on appropriately drained
constant topsoil depth. land with previously poor drainage. Costs
can be relatively high, including the expense
of materials, drain installation and engineer-
11.3.2.2 Contouring
ing design. However, there is considerable
Where water runoff and soil erosion con- information available relating to the depth of
cerns are great, orchards can be established installation, spacing, size and type of drains,
parallel to contours, so that orchard opera- as well as design and layout procedures
tions can be undertaken as nearly as practi- from standard soil-engineering texts
cal on the contour (Plate 11.4). Such planting (Schwab et al., 1966) and, in many regions,
systems were originally designed so that from companies specializing in soil drainage.
tillage operations could be undertaken
across the slope gradient, reducing the veloc-
ity and erosive power of overland flow. The 11.3.3 pH adjustment
decline in orchard cultivation following the
advent of herbicides has greatly lessened If soil pH is not optimum for apple tree
tillage operations in orchards. However, growth, the best time to correct the problem
planting orchards on the contour allows is prior to tree planting when ameliorative
orchardists to subsequently establish strip amendments can be easily worked into the
cropping systems, where vegetation-free, in- soil without concern for damaging the newly
row herbicide strips alternate with between- planted trees.
Excessively low pH values are more com- neutralizing ability and possibility of conta-
monly encountered and are corrected via mination with toxic elements. Packing-
lime applications. There are a range of suit- house lime (originally calcium hydroxide)
able liming materials whose ability by weight can be an economical source of locally avail-
to neutralize acidity varies and is often able liming material, but probably requires
expressed as a percentage of pure calcium regrinding to achieve effectiveness, since it
carbonate (defined as 100%) (Table 11.2). often forms large insoluble granules after
Limes are usually comprised primarily of absorbing water and carbon dioxide.
calcium compounds but are applied to neu- Lime is most effective when finely ground
tralize acidity rather than for their direct because its reaction rate depends on its sur-
contribution to the calcium nutrition of face area in contact with the soil. The reac-
apple trees. Magnesium-containing limes, tion is also speeded by intimate contact with
such as dolomite, should be considered the soil, which may be achieved by plough-
when soil magnesium contents are low and ing and discing the soils after surface appli-
cultivars such as ‘McIntosh’ are being cations. Little change in bulk soil acidity was
planted that are susceptible to the develop- measurable several years after rotavating
ment of magnesium deficiency in heavy coarse fragments of packing-house lime into
crop years. Some liming materials, such as the soil profile. Without incorporation, pene-
calcium oxide or calcium hydroxide, neu- tration of even finely ground lime can be
tralize acidity more rapidly. These materials exceedingly slow, as indicated by significant
can be difficult to apply, since calcium oxide pH changes only being detected in the sur-
can form larger, less soluble granules after face 10 cm 5 years after application at 6 t
absorption of water from the soil. Calcium ha1 of calcium hydroxide to the soil surface
hydroxide can be difficult to spread as a fine without incorporation (Neilsen et al., 1981;
powder, especially on windy days. Fig. 11.7).
Sometimes, inexpensive, locally available The amount of lime to apply varies pri-
sources of liming material, including marl, marily with the texture and the organic-mat-
basic slag, fly ash and packing-house lime, ter content of the soil. It is usually calculated
are available to the orchardist. Their rela- as the amount required to achieve target pH
tively cheap cost must be balanced by con- values to a fixed soil depth (usually 20 cm)
sideration of their effectiveness, actual by equilibrating a representative sample of
Table 11.2. Commonly used liming materials and their neutralizing value (data adapted from Tisdale et
al., 1993).
Molecular Neutralizing
weight valuea
Material Common name(s) (g mol1) (%)
Calcium oxide Quicklime, unslaked lime, 56 179

CaO burned lime
Calcium hydroxide Hydrated lime, slaked lime, 72 136
Ca(OH)2 builder’s lime
Calcium–magnesium carbonate Dolomite 184 109
(Ca, Mg)CO3
Calcium carbonate Agricultural limestone 100 100
CaCO3
Calcium silicate Basic slag 116 86
CaSiO3
aAs a % of pure calcium carbonate set as 100%. To be equivalent to a given quantity of calcium
carbonate, neutralizing values exceeding 100% required proportionately smaller amounts of material.
10
Depth (cm)
20
30
40
50
60
5 10 15 20 4 5 6
Soil exchangeable Mn pH
Fig. 11.7. Surface application of various calcium compounds to a ‘Spartan’ apple orchard in November,
1994, followed by measurements of average soil exchangeable manganese (Mn) and pH with depth for each
of the 6000 kg ha1 Ca(OH)2(––), 12,000 kg ha1 CaSO4 (.....) and control (––) treatments in 1979.
Statistically significant differences indicated at the 0.01 (**) and 0.05 (*) levels.
the soil in a buffer solution. The Shoemaker, orchard soils, pH values ranging from 3.9 to
McLean and Pratt (SMP) single-buffer proce- 4.4 have been measured in the tree row rela-
dure has been widely used to make these cal- tive to mid-alley values of 5.4–6.7 (Ross et al.,
culations by many soil-testing laboratories 1985). With the possibility of such large pH
(van Lierop, 1990). When soil pH is uni- differences on old orchard land, it may be
formly low in an orchard block, a uniform appropriate to apply differential, higher
rate of lime can be applied. However, consid- rates of lime in old tree rows.
erable pH variation can occur due to previ- Subsoil pH can be important, since apple
ous management practices, including roots when planted must begin to proliferate
nitrogen fertilization, irrigation and herbi- and grow at depths at and below 30 cm. It
cide use in land previously in orchard. For can be difficult to mechanically incorporate
example, zones of low pH near previous lime effectively to these depths. In regions
apple tree locations where acid-forming where very acid subsoils (pH < 5) exist,
nitrogen fertilizer applications were concen- improved plant growth has been observed
trated have been reported in Washington several years after incorporation of gypsum,
State orchards (Benson, 1968). In coarse-tex- which has acted by reducing soil concentra-
tured, low-organic-matter, sandy-loam tions of toxic aluminium (Sumner et al., 1986).
There is also some potential to acidify The use of organic materials as soil
soils with excessively high pH values. For amendments usually involves surface appli-
example, the pH of calcareous soils can be cation to the proposed tree row and rotavat-
reduced by application of acidifying coming to mix materials throughout the root
pounds, which also increase the availability zone prior to planting. Effective application
of other nutrients by neutralizing calcium rates are frequently in the 30–60 t ha1 range
carbonate. Acidifying materials include ele- since 10–20% by volume of material must be
mental sulphur, sulphuric acid, aluminium added in order to meaningfully alter bulk
sulphate and ammonium polysulphide. soil properties. The nutrients co-applied with
Elemental sulphur is often the most effective the organic materials can improve long-term
of soil acidic substances and research on soil fertility. However, organic materials
orchard soils has indicated that effectiveness, must be well mixed in the soil to avoid creat-
as with liming, increases with soil incorpora- ing excessively saline conditions, since many
tion and finer particle sizes (Neilsen et al., organics have high salt readings. There may
1993). Complete neutralization of calcium also be difficulties in regulating the nitrogen
carbonate is often too expensive, due to the available to the trees, since applied organics
large quantities of calcium carbonate in can act to either release or immobilize nitro-
many soils. Some success has been achieved gen depending upon the carbon : nitrogen
by acidifying a portion of the root zone by ratio of the materials, as well as temperature
band application of high rates of acidifying and moisture conditions in the soil environ-
fertilizers, such as ammonium thiosulphates ment.
or polyphosphates, after planting. It can be It is also possible to make applications of
extremely difficult to reduce the pH of saline these various organic materials to the surface
soils, unless there exists the possibility of
of soils around trees immediately after plant-
leaching large quantities of salt from the soil
ing (Fig. 11.8). Such mulch applications offer
profile via irrigation. A wiser strategy for
several benefits, including moderation of soil
such situations may be to avoid planting
temperature extremes, conservation of soil
such sites in the first place.
moisture, reduction in weed competition,
improved nutrient availability (especially
11.3.4 Other soil amendments phosphorus and potassium) and long-term
improvement in the aggregation and perme-
In addition to pre-plant lime and fertilizer ability of poorly structured soils. A current
applications to improve the performance of area of research involves the suppressive
a newly planted apple orchard, there are sit- effects the addition of organic materials have
uations where applications of other soil upon soil-borne pathogens, including
amendments may be advantageous. Phytophthora root rot of apples. An important
Recently, the use of various organic amend- concern in many regions is that mulches can
ments and mulches has received renewed provide an excellent habitat for orchard
attention, due to concerns about the nega- voles (mice). These animals can devastate an
tive impacts that conventional orchard pro- apple orchard by girdling the trunk and/or
duction practices are having on soil quality the roots. Where voles are a concern, a vege-
(Glover et al., 2000). Such applications often tation-free strip along the tree row is usually
depend upon the ready availability of eco- a better choice, as it tends to discourage
nomical sources of suitable organic materi- rather than encourage orchard mice.
als. Locally available sources can reduce Effective use of organic materials as
transportation costs for such materials, amendments or mulches to improve short-
which are often wet and bulky. Source mate- term apple tree vigour and production are
rials may include various hays and straw, not guaranteed but are more likely on sites
grape pomace, manures or sewage biosolids with coarse-textured, low-organic-matter,
and, increasingly, composted combinations shallow soils where otherwise limiting nutri-
of various of these materials. ent and water stresses may occur.
individual biotic factors, there is no consensus

across all fruit-growing regions concerning
the most important single factor or whether
several factors act simultaneously or sequen-
tially or interact with adverse environmental
conditions to weaken growth of replanted
apple trees.
11.4.1 Fumigation
Application of broad-spectrum chemical bio-

cides to old orchard soils prior to replanting
has, however, often been effective in improv-
ing the initial growth and subsequent yield of
replanted apples, despite uncertainty con-
cerning the exact causal factor(s). For exam-
ple, large yield increases were observed for
‘Royal Gala’ apple on M.7a rootstock after
replant application of Metham-sodium at var-
ious rates in Washington State. Improved
growth has also been reported after use of
other general fumigants in various fruit-
growing regions (Table 11.3). Local research is
Fig. 11.8. Black plastic, lucerne and paper mulches in
a high-density ‘Spartan’ apple orchard in autumn
probably the most valuable guide to effective
1998. fumigation, due to the apparently site-specific
nature of the problem in different regions.
Several general practices, however, apply
11.4 Replant-site Preparation to effective fumigation, including: good site
preparation, which usually means removal of
An important consideration when replanting old trees and stumps, with as many roots as
a site where apples have previously grown is possible; mitigation of other soil limitations
the possibility of inhibited growth of the via application of required fertilizers, lime or
newly planted apple trees, which has been other soil amendments, followed by plough-
observed in most major apple-growing areas ing, discing and deep ripping of soil, as
and attributed to the consequence of growing appropriate; and application of the fumigant
the same crop repeatedly on the same soil. It to warm (10–18°C) and moist soil. These lat-
is likely that the occurrence of such problems ter conditions often occur in either spring or
will increase as more planting occurs on land autumn. Fumigation in the autumn has the
previously in orchard and as planting systems advantage of dissipation of the fumigant
are established with the intention of more fre- from the soil prior to the planting of young
quent renewal with new cultivars. Poor trees. Spring fumigation can be carried out,
growth of apples on replant sites is sometimes but caution is required, since soil containing
referred to as specific apple-replant disease residual fumigant can damage young trees.
(SARD) but is not always specific, having The dissipation of fumigants from the soil is
been observed on sites not previously having affected by temperature, being more rapid on
apples (Sewell, 1979). Several causal factors warm soils. A germination test can be carried
have been identified, including a wide range out to determine whether all fumigant has
of pathogenic organisms, such as various dissipated from the root zone prior to plant-
species of actinomycetes (Otto et al., 1994), ing in the spring. This test compares the ger-
fungi and nematodes (Yadava and Doud, mination of seeds of either cress, lettuce or
1980). Despite the identification of numerous radish applied to the surface of fumigated
Table 11.3. Fumigation treatments successfully used to overcome replant problems in old orchard soils.
Effective field application

Compound rate Considerations
Chloropicrin 25–50 ml m2 Commercial application recommended

(trichloromethane) due to high toxicity; apply within proposed
row, seal with plastic; similarly effective
3–18°C
Formalin 400–600 ml m2, Apply to moist soils with water
(formaldehyde) apply with 2–4 l m2 of water
Vapam (Metham-sodium) 90 ml m2 Apply to wet soil; can be applied
(sodium N-methlydithio- with irrigation water
carbamate)
Basamid (98% dazomet) 30–50 g m2 Granular, releases toxic gas upon soil
(tetro-hydro-3,5-dimethyl-1,3,5- contact; requires incorporation to target
thiadiazine-2-thione) depth; seal with water, preferably plastic
and untreated samples of the moist soil 11.4.2 Alternatives to fumigation

sealed in small glass jars and placed in warm,
sunny conditions. The test can be repeated at Fumigation can be expensive, difficult to
2-day intervals until germination occurs and accomplish safely and detrimental to bene-
the date of tree planting can be established. ficial soil microorganisms, such as mycor-
The use of broad-spectrum fumigants is rhizae. Consequently, environmentally
non-discriminatory and can decrease popu- benign alternatives are being continuously
lations of potentially beneficial microorgan- sought, although none has yet achieved the
isms, such as vesicular arbuscular wide-ranging success associated with fumi-
mycorrhizae, which have a role in the uptake
gation. For example, planting of marigold
of plant nutrients such as phosphorus. This
(Tagetes patula L.) as a cover crop has sup-
may be the reason why the application of rel-
pressed root-lesion nematodes (Pratylenchus
atively high rates of phophorus, especially in
spp.) in New York, where nematodes have
the mono- and diammonium phosphate
been judged an important component of the
forms, within the tree planting hole in fumi-
replant-disease problem (Merwin and Stiles,
gated, pasteurized or fungicide-treated soils
1989). Similarly promising suppressions in
has ensured vigorous growth of young apple
trees in old orchard soil (Slykhuis and Li, the American dagger nematode (Xiphinema
1985; Neilsen and Yorston, 1991). americanium) populations have been
Not all soils will respond to fumigation observed after planting certain Brassica
and, in some apple-growing areas, a seedling plants (Halbrendt and Jing, 1994).
bioassay to test for the presence of apple- Replacement of disease-infested soil with
replant disease has been developed and is new soil in the planting hole is possible, but
offered by some soil-testing laboratories. it can be difficult to locate sufficient high-
Tests usually involve comparisons of the quality replacement soil and long-term
growth of apple seedlings in untreated or results can be disappointing, as newly
fumigated soils after 12–14 weeks’ growth in planted trees soon grow into the original
pots under greenhouse conditions. soil. First-year tree growth has, however,
Fumigation is recommended when large been increased on replant sites by planting
growth increases (> 150%) are measured for hole amendments involving applications of
seedlings grown in treated soil (Sewell et al., non-orchard soil (Peryea and Covey, 1989)
1988). Seedling bioassay tests can thus pro- and various organic mixes, often in associa-
vide valuable information on whether to tion with high rates of monoammonium
fumigate or not. phosphate fertilizer (Neilsen et al., 1994).
11.5 Orchard Planning 11.5.1.3 Organic apples

The production of organically grown apples
11.5.1 Cultivar selection has expanded in recent years, but although
this trend continues, the organic part of the
Projections for the world apple situation industry remains a very small part of the
indicate that growers will continue to face a total (see Chapter 22).
period during which production will exceed
demand (see also Chapter 2). Cultivar selec-
tion is one of the most critical decisions 11.5.1.4 Direct marketing/pick your own
made as new orchards are planned. The grower who is able to market fruit
Although many factors must be considered directly to the consumer through one of
in selecting cultivars, the planned marketing many types of retail outlets or a pick-your-
strategy is of utmost importance. In this sec- own operation has a real advantage. The
tion, we shall introduce the subject of culti- wholesale grower has essentially no contact
var selection; more detailed coverage is with the ultimate consumer, but rather must
presented in Chapters 2, 4, 11 and 24. deal with the whims of a chain-store buyer.
In contrast, in a direct marketing operation,
it is quite feasible to ‘educate’ the customer
11.5.1.1 Wholesale fresh market by offering samples and thereby to sell more
Today, a typical supermarket in the USA will and different cultivars. It is not unusual for a
display 12–15 different cultivars of apples, direct-market grower to offer 30–40 cultivars,
several of which are of relatively recent often including old or ‘heritage’ cultivars,
introduction. Growers are scrambling to find particularly those of a regional nature, which
the newest cultivar for which they hope to are not available in grocery stores.
get the early peak price usually received as a
successful new cultivar first comes on the
market. Unfortunately, in the rush to ‘get 11.5.2 Rootstock selection
there first’, growers sometimes plant rela-
tively untested cultivars, which prove to be In sharp contrast to growers of other tree
fruits, apple growers are very fortunate in
unprofitable because of poor adaptation to
having a wide array of dwarfing rootstocks
their climate, serious cultural problems or
available (see Chapter 5). The major factors
lack of sufficient production for the cultivar
that have an impact on rootstock selection
to be accepted in the wholesale market.
include: vigour, precocity, yield, yield effi-
Given its many facets, cultivar selection
ciency, disease and insect resistance, anchor-
remains one of the most vexing issues facing
age, fruit size and quality, suckering and, last
the fresh-market apple grower. but not least, grower experience and man-
agement skills.
11.5.1.2 Processing market
In the processing market, the cultivar picture 11.5.2.1 Vigour
is much more stable than in the wholesale There is ample evidence that, in the early years
fresh market. For utilization as tinned or of an orchard, yield per unit area is largely
frozen apple slices or apple sauce, specific dependent on the number of trees per hectare
cultivars are preferred and a premium price (see Chapter 15). This consideration is one of
is paid for them. Although the cultivars of the driving forces in the movement to increas-
choice vary with locality, the market is rela- ingly dwarfing rootstocks and intensive train-
tively stable. Unfortunately, however, the ing systems. The key to successful orchard
depression of prices related to overproduc- design is to plant trees close enough to obtain
tion and worldwide competition is of con- good early yields while avoiding future tree-
cern, particularly with the apple-juice management problems and not sacrificing fruit
market (see Chapters 2 and 24). yield and/or quality as the trees mature.
The wide range of rootstock vigour avail- British Columbia (Quamme et al., 1997), the
able to today’s apple grower is nothing less two high-density systems outyielded the
than remarkable. For example, in the 10-year lower-density systems by three to five times
NC-140 rootstock trial, completed in 1989 over the first 4 years. It must also be realized,
(NC-140, 1991), tree size, as indicated by however, that, compared with traditional
trunk cross-sectional area (TCA), varied by a low-density orchards, intensive orchards
factor of about 17 – from 10 cm2 for trees on have higher establishment costs and require
M.27EMLA to 172 cm2 for trees on MAC.24 greater management skills.
(Fig. 11.9). As the young orchard develops, overall
canopy volume continues to be the overrid-
ing factor influencing yields (Plate 11.5). For
11.5.2.2 Precocity
this reason, training systems that encourage
A very notable characteristic of the dwarf early canopy development tend to maxi-
rootstocks is their remarkable ability to mize yields. For example, in the NC-140
induce precocity in scion cultivars. In fact, it orchard systems trial, trees trained to the
is often a concern that such trees tend to vertical axis had higher early yields per tree
flower excessively in the first year or two fol- and per hectare than trees trained as either
lowing establishment in the orchard. The a slender spindle or a central leader (Barritt
potential problem is the tendency for some et al., 1997b). Part of this difference is proba-
rootstocks, such as Mark, to ‘runt out’ (essen- bly due to the minimal pruning of the verti-
tially stop vegetative growth) as a result of cal axis compared with the other two
excessively heavy fruiting in the second or systems.
third year.
11.5.2.4 Yield efficiency
11.5.2.3 Yield
Yield efficiency is calculated by dividing the
The interest in high-density orchards has weight of fruit produced per tree by the
been driven by the desire for early returns on TCA. In essence, yield efficiency is the ratio
the orchard investment, and dwarfing root- of fruit production to wood production. In
stocks are mandatory for their precocity as general, dwarf rootstocks have considerably
well as their vigour control. In a study in higher yield efficiencies than the more vigor-
Fig. 11.9. Cross-sections of trunks of 10-year-old ‘Starkspur Supreme Delicious’ on rootstocks ranging from
the vigorous MAC.24 to the very dwarfing M.27EMLA.
ous rootstocks. There is, however, a point of 11.5.2.7 Fruit size, maturity and quality
diminishing returns where the tree produces
Over the past 20 years, several researchers
so much fruit and so little wood that tree
have evaluated the influence of apple root-
canopy size is seriously restricted. Thus, the
stocks on apple maturity, size, quality and
allocated space is not fully occupied and the
storage life. Although differences in fruit size
yield per hectare does not meet expectations.
have not always been consistent, trees on
For example, in some trials, trees on Mark
OAR1, P.1 and M.27 have tended to produce
have had very high yield efficiencies, but
smaller fruit than trees on M.9, M.26 and B.9
have produced such small canopies that total
(NC-140, 1991; Barritt et al., 1997a).
yields per tree and per hectare are often
lower than for trees on M.9. Fortunately, none of the three that are prone
to produce small-sized fruit are among those
being widely planted. Rootstock effects on
11.5.2.5 Disease and insect resistance maturity, quality and storage life have been
In making a rootstock selection, disease sus- relatively small in studies over multiple sites
ceptibility is an important consideration, and years and therefore need not be a major
because of the tremendous impact that cer- concern in making rootstock choices (Autio
tain diseases can have (see Chapter 18). et al., 1991).
Whole blocks of trees can be devastated by a
major outbreak of fire blight (Erwinia 11.5.2.8 Suckering
amylovora (Burrill) Winslow et al.).
Unfortunately, almost all of the dwarfing With most apple rootstocks in use today,
rootstocks being planted are highly suscepti- suckering has not been a serious concern.
ble to fire blight, as are many of the newer One notable exception is the semi-dwarf
scion cultivars. A fruit grower needs to be M.7, which is well known to sucker more
particularly alert when both the scion and than others. When numerous, there is the
rootstock are highly fire blight-susceptible obvious labour cost for annual removal, but
and extra care must be devoted to minimiz- there is an additional concern when the root-
ing the likelihood of an outbreak. Viruses stock is highly susceptible to fire blight.
and soil-borne fungi can also be trouble-
some. There is little in the way of insect resis-
11.5.2.9 Grower experience/management
tance in apple rootstocks (see Chapter 19). A
skills
particularly well-known exception to this
statement is the resistance to woolly apple Most growers have been making the transi-
aphids (WAA), which was incorporated in tion from orchards on seedling rootstocks at
the Malling–Merton rootstocks by English 100 trees ha1 to those on semi-dwarfing
researchers who crossed Malling rootstocks rootstocks at 300–450 trees ha1. It is much
with ‘Northern Spy’, which has excellent easier to go to a high-density orchard of
WAA resistance. 1000–1500 trees ha1 from a semi-dwarf
orchard of 450 trees ha1 than from a con-
ventional orchard of 100 trees ha1.
11.5.2.6 Anchorage
As new rootstock candidates become
Among the rootstocks available today there available, growers should test them in their
is the complete spectrum from those that are own orchards, but relatively untested root-
completely self-supporting (MM.111) stocks should be planted only in small num-
through to those that have a tendency to lean bers until their suitability is established.
(M.7) and to those for which trunk support is Unfortunately, new rootstocks are sometimes
mandatory (M.9). This categorization is made available to commercial growers
based on whether or not the rootstock is able almost as soon as they are available to
to support the fully fruiting tree in an researchers. It takes a minimum of 5–10
upright position without additional support years to assess the strengths and weaknesses
(see Chapter 5). of new rootstocks. In the meantime, the
planting of ‘known quantities’, such as one selection of cultivars and rootstocks has
of the M.9 clones, is a better choice. been even further complicated by the explo-
A mistake made by some growers has sion of choices of new strains of both scion
been to take information from a different cultivars and rootstocks. For example, there
part of the country or world and to attempt is a seemingly ever-changing array of
to duplicate a particular orchard design and strains of ‘Gala’ and many different clones
system on their own site without consider- of M.9 rootstocks. Although individual
ing the differences in soil, climate and man- nurseries tend to concentrate on a limited
agement skills. For example, a particular number of choices, it is not surprising that
tree spacing in a northern climate, such as errors can and do occur. The best insurance
western New York or Michigan, may be against disappointment is to deal with a
much too close in the mid-Atlantic area, reputable nursery.
where tree vigour is notably greater. This
has become apparent in data from regional
11.6.1.1 Feathered trees
trials, such as the NC-140 rootstock trials,
over the past two decades (NC-140, 1991; With the trend towards more intensive
1996), which show that, in general, trees in orchards, productivity in the early years has
areas with longer growing seasons tend to become critical to economic success. One
be larger. way to encourage early fruiting is to plant
feathered (branched) trees, which can save a
year in getting trees into production (see
11.5.3 Row orientation Chapters 6 and 14).
Since the advent of hedgerow plantings,

11.6.1.2 Planning
researchers have sought to ascertain the ideal
way to orientate rows to maximize the inter- The lack of adequate planning sometimes
ception of light. Factors that interact and leads to serious mistakes in buying trees. If a
therefore complicate this issue include lati- grower decides at the last minute to order
tude, time of year, time of day, tree height trees (for delivery in a matter of weeks or
and shape and row spacing. Results from even months), he/she is frequently left with
several researchers indicate that, in general, a minimal choices of specific cultivar/root-
north–south row orientation provides both stock combinations. This is particularly true
better light interception and a more even dis- if the combination of choice is in high
tribution throughout the tree canopy than an demand by others. Settling for one’s third or
east–west row orientation and is therefore fourth choice is very often a serious error in
preferred (Cain, 1972; Jackson, 1980). judgement. It is far better to order exactly the
cultivar/rootstock desired, preferably 2
years in advance, to ensure its availability.
11.6 Tree Establishment Nurseries often offer a discount for trees
ordered well in advance.
11.6.1 Tree quality
11.6.1.3 Tree condition and arrival
In the establishment of a new orchard, tree
quality has to be a top priority. An orchard Trees must arrive in prime condition and on
is a long-term commitment and success is, time. Trees that have either dried out or been
to a sizeable degree, dependent upon the frozen during transit should be rejected,
quality of the trees planted. First among the because they will probably grow poorly, if at
tree-quality attributes has to be trueness to all. Likewise, trees showing signs of either
name for both the scion cultivar and the disease infection or insect infestation should
rootstock. There is a tremendous array of be rejected. It is wiser to wait an additional
both cultivars and rootstocks propagated year if suitable replacement trees are not
and sold by nurseries. In recent years the available.
11.6.1.4 Tree storage

When the trees arrive they must be kept dor-
mant and the roots moist. Cold storage is fre-
quently required, but great care must be
taken to ensure that the trees are not exposed
to ethylene. A cold-storage room either con-
taining apples or in which apples were
stored will contain a sufficient concentration
of ethylene to kill or severely injure apple
trees. The room used to store trees must not
contain apples and, if the room has previ-
ously stored apples, the atmosphere must be
flushed for a lengthy period to minimize eth-
ylene concentration. If suitable refrigerated
storage space is unavailable, trees can be
‘heeled in’ in soil or sawdust on the shaded
side of buildings for short-term holding.
11.6.2 Tree planting
11.6.2.1 Season of planting

In regions without excessively cold winters,
there are advantages to planting trees in the
autumn. Although autumn-planted trees will
make no shoot growth until spring, roots
will grow when soil temperatures are
approximately 5°C or higher. These addi-
tional roots can be of considerable value to
the tree when shoot growth is initiated in the
spring. Drawbacks to autumn planting
include lack of tree availability, conflict with
harvest, potential damage by voles, rabbits
and other wildlife during the winter (Fig.
11.10) and potential winter injury. In irri-
gated regions, dry soils in the early spring
may make it difficult to get good root growth
on autumn-planted trees. In areas that expe-
rience frequent freezing and thawing cycles, Fig. 11.10. A young apple tree that suffered
‘frost heaving’ can push newly planted trees girdling injury from both mice and rabbits. The
upwards, exposing the upper roots. Such injury occurred while snow covered the orchard.
trees must either be replanted or have soil Mice girdled the base of the tree under the snow
brought in to cover the roots. and the rabbits, able to travel on an ice coating on
the snow, ate bark as high as they could reach.
It is well documented in humid areas that
Uninjured bark is apparent in the area in between,
trees planted in late winter to early spring which was snow-covered.
grow far better than those planted later in
spring. Spring weather is notoriously variable
and wet periods that delay planting are fre- appropriate soil conditions. In arid areas, dry
quent. It is critical that trees be available on soils in the spring may necessitate some delay
site so that they can be planted during the in spring planting or at least require available
earliest suitable weather associated with water to adequately moisten planting-hole
soil. Whether the trees are planted in autumn Regardless of the technique and equipment
or spring, there must be adequate, but not used, certain standards must be met. The
excessive, soil moisture present at planting most critical of these is the location of
and during the ensuing months. the bud union. After the tree has settled and
the soil in the planting hole has consolidated,
the bud union should be 4–6 cm above the
11.6.2.2 Tree spacing and arrangement
soil surface; this means that when initially
For many decades, apple trees have been planted, the bud union should be 6–8 cm
planted closer in the row than between rows; above the soil surface. If the union is buried
thus they can be said to be in a rectangular or settles below the soil surface, scion root-
design as opposed to trees being planted ing can overwhelm the dwarfing characteris-
equidistant in both directions (so-called tics of the rootstock, leading to excessive tree
square design). Under the overall umbrella vigour and tree crowding. If too much root-
of rectangular designs are an amazingly stock is left exposed above the soil, burr-
broad spectrum of orchard designs, which knots (masses of root initials) may develop.
are described in Chapter 15. These burr-knots lead to deformed trunks
(Fig. 11.11), can serve as entry points for fire
blight bacteria and for insects such as the
11.6.2.3 Placement of pollenizers
dogwood borer and can also lead to exces-
Almost all cultivars of apple require cross- sive dwarfing. In the 1980s, a recommenda-
pollination to set commercial crops, and tion was made that trees be budded higher
there have been many different approaches in the nursery (by 15–20 cm) so that trees
taken to meet these pollination needs. With could be planted deeper, theoretically offer-
the widespread utilization of the hedgerow ing additional tree support, especially for
design, a commonly accepted practice is to trees on M.7, which tend to lean.
place pollenizer trees (often flowering crab Unfortunately, high-budded trees set in clay
apples) about every 15 m in each row and to soils did not thrive (Lyons et al., 1983) and
offset the pollenizer trees in adjacent rows. the practice was largely abandoned.
Particularly with very dwarfing rootstocks, The regulation of planting depth is rela-
no space is allocated to the crab apple poll- tively easy with trees planted by hand, but
enizer trees. They are set between adjacent can be more of a challenge when using a tree
fruiting trees and are trained to a tall cylin- planter. It may be necessary to have workers
drical shape. Since crab apples flower so pro- follow behind a tree planter to adjust plant-
fusely, the severe pruning that is required ing depth of individual trees. Such adjust-
does not restrict flowering. Among the ment is far easier immediately after planting
advantages of crab apples as pollenizers are than after the soil has settled around the
their annual flowering without the need of roots.
thinning, no mix-up of fruit at harvest and a
wide choice of flowering time to match most
cultivars. A common practice is to plant two 11.6.3 Tree support
to three different crab apple cultivars to bet-
ter span the flowering period of the main For the last several decades there has been
cultivar. Attention must be paid to selecting an ever-accelerating trend towards the use
crab apple cultivars with resistance or immu- of size-controlling rootstocks in new apple
nity to fire blight. orchards; few apple trees are planted today
on vigorous seedling or even the relatively
vigorous MM.111 rootstocks. Another part
11.6.2.4 Planting depth
of this trend has been a shift to increasingly
In planting apple trees, there has been a pro- dwarfing rootstocks. In the opinion of most
gression from doing so by hand with a people, the more dwarfing stocks (M.26 size
shovel to the use of augers to drill holes to or smaller) require support because of the
the use of mechanical tree planters. inability of such trees to support heavy
high costs, there is a temptation to seek

cheaper approaches by using fewer or smaller
posts or by forgoing the wire to support the
wooden or metal stakes. When the trees are
1–2 years old and largely vegetative, a mini-
mal support system, such as an unsupported
conduit, appears to be sufficient; by the fourth
or fifth year, however, it becomes apparent
that a minimal system is totally inadequate.
As a rule, it is far easier and cheaper to install
an adequate support system at the outset,
rather than to have to replace an inadequate
system that fails when the trees begin to bear
heavily. It is very easy to underestimate the
amount of stress imposed upon a support sys-
tem by apple trees that have a full crop, expe-
rience a heavy rainfall event and are then
buffeted by high winds a few days or even
weeks before harvest.
Some growers avoid the necessity for tree
support by using severe pruning practices,
basically following the procedures recom-
mended for central-leader trees by Heinicke
(1975). If the leader is headed by one-half
each year, it may be possible to grow trees on
a rootstock such as M.26 without support.
Fig. 11.11. The lower trunk of a 10-year-old There are data, however, which show that
‘Starkspur Supreme Delicious’ on M.26EMLA. Just such severe pruning reduces yields in the
below the bud union is a large burr-knot (group of early years of the orchard, probably by
root initials), which has deformed the normal shape enough to more than pay for a support,
of the trunk and is a potential entry point for fire which would avoid the need for such heavy
blight bacteria, as well as certain insects, such as pruning (Barden and Marini, 1998).
the dogwood borer.
11.6.4 Pruning at planting

crops (see Chapter 15). Support can range
from a short post providing only trunk sup- Prior to the introduction of feathered trees,
port to 3 m stakes, which are in turn sup- most apple trees were planted as ‘whips’,
ported by a wire (Fig. 11.12). There are which were unbranched trees ranging in
intermediate types of supports, including height from about 1.5 to 2 m. Such trees were
single posts of metal or wood and wire trel- routinely headed at a height of 70–100 cm.
lises, as well as elaborate wood and wire or Such heading was to re-establish the balance
steel and wire trellises. Each has its special between the top of the tree and the root sys-
attributes and drawbacks and the choice is tem, which had been damaged by being dug
difficult. The basic question for trees on and shipped bare-root, as well as to induce
dwarfing rootstocks is not ‘Should the tree buds to break and to produce shoots that
be supported?’ but rather ‘How should the would eventually become the lower tier of
trees be supported?’ scaffold limbs.
One of the greatest concerns with any type With the availability of well-feathered
of tree support is the relatively high cost, both trees (multiple branches at the desired height
for the materials and for the labour for instal- and angles), the need to head the newly
lation (Fraser and Oakes, 1999). Because of the planted tree to induce branching was no
Fig. 11.12. Young vertical axis apple trees in southern British Columbia. Individual trees are supported by
bamboo poles, which are in turn supported by a wire at approximately 2.5 m.
longer valid. Various recommendations have but it must be kept in mind that we must first
been made for such trees, including no head- grow the trees, and good growth during the
ing at all if the feathers are acceptable in both first year is vital.
number and location. Others suggest heading
at 25–30 cm above the top feather. Opinions
also vary as to whether the feathers should 11.7 Summary and Conclusions
be headed. The issue here is the balance
between the root system and the top of the The establishment of apple orchards involves
tree. As discussed above, if trees are planted a great many decisions, each of which has a
in the autumn or very early spring, root long-term impact on the success of the opera-
growth can precede bud break and therefore tion. Climatic considerations are of para-
may meet the water needs of the new mount importance and encompass both
growth. If, however, the buds break soon broad geographical and very local factors.
after planting, the tree may suffer from lack Apple trees require approximately 1000 h of
of water, will make little or no growth and chilling during their dormant season and
may even die from desiccation. In the arid perform best where winters are not extremely
north-western apple areas of North America, cold and summers not excessively hot. The
there is a tendency for shoot growth to flowers and young fruit of apple are subject
slightly precede root growth, which seems to to spring-frost injury, so site selection is based
create a sensitivity to water stress if moisture to a sizeable degree on this consideration.
conditions are not optimum. On the basis of Locations on the leeward side of lakes or
the available information, it is our recom- other large bodies of water are excellent
mendation that the leader and feathers be because of the temperature-moderating effect
headed unless the trees were planted in the of the water. Other preferred sites are those
previous autumn. The goal is to get new that have good air drainage due to slope or
orchards into production as soon as possible, position above the surrounding area.
There are likewise many aspects of soil The planning of a new orchard should
that are vitally important in choosing a site start at least 2 years prior to planting to allow
for an apple orchard. Among the physical the ordering of the desired cultivar/rootstock
characteristics that must be considered are combination(s). There is an increasingly broad
soil texture, structure and depth, because of list of potential cultivars, and they vary not
effects on drainage, nutrient availability, only with the climate but with the market to
water relations, aeration and equipment be served. Likewise, there are many root-
travel. Because of slow movement in the soil, stocks from which to choose, but the world-
lime to adjust soil pH as well as applications wide trend is towards increasingly dwarfing
of phosphorus and potassium should be rootstocks. Whatever the choice of cultivar
made prior to planting so that they may be and rootstock, the trees should come from a
ploughed down into the root zone. Other reputable nursery, be planted as early as pos-
potentially useful procedures may include sible in the spring and be supplied with ade-
breaking up impermeable layers in the soil, quate moisture. There are many options as to
levelling, contouring, terracing, improving tree spacing and arrangement, initial pruning,
drainage or adding additional soil amend- tree support and training systems. Regardless
ments. In situations where a replant problem of the cultivar, rootstock and orchard system
exists, there may be a need for fumigation; selected, the success of the orchard will
efforts are under way to develop more envi- depend to a large degree on the characteristics
ronmentally benign alternatives. of the site on which it is planted.
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12 Nutritional Requirements of Apple
Gerry H. Neilsen and Denise Neilsen

Agriculture and Agri-Food Canada, Pacific Agri-Food Research Centre, Summerland,
British Columbia, Canada
12.1 Nutrient Requirements 267

12.2 Direct Determination of Apple-tree Nutrient Status 268
12.2.1 Leaf nutrient concentration 268
12.2.2 Fruit nutrient concentrations 270
12.3 Nutritional Implications of Rooting Characteristics 271
12.4 Soil Testing 273
12.5 Augmenting Apple Nutrition 274
12.5.1 Soil applications 274
12.5.2 Foliar applications 276
12.6 Special Nutrient Considerations 277
12.6.1 Organic production 277
12.6.2 Integrated fruit production 278
12.7 Individual Nutrients 278
12.7.1 Nitrogen 278
12.7.2 Phosphorus 286
12.7.3 Potassium 288
12.7.4 Magnesium 289
12.7.5 Calcium 290
12.7.6 Sulphur 292
12.7.7 Boron 293
12.7.8 Zinc 295
12.7.9 Iron 296
12.7.10 Manganese 297
12.7.11 Copper 298
12.1 Nutrient Requirements centrations within the plant include nitro-

gen, phosphorus, sulphur, potassium, cal-
In common with many plants, apple trees cium and magnesium. Minerals usually
require 16 elements for successful comple- comprising lower (p.p.m.) concentrations
tion of their life cycle. Among these ele- include iron, manganese, copper, zinc,
ments are carbon, hydrogen and oxygen, boron, molybdenum and chlorine. It is diffi-
which are important non-mineral elements cult to calculate the total nutrient require-
and major constituents of organic materials. ments for apple trees, since it is necessary to
Mineral elements contained in high (%) con- account for nutrients contained in the peren-

268 G.H. Neilsen and D. Neilsen
nial framework of the trunk and roots as fruit quality, some emphasis has been placed
well as those contained in leaves, new upon use of fruit nutrient concentrations
shoots and roots, which are produced annu- (especially calcium) to optimize the harvest
ally. Some measurements have been under- and storage quality of fruit.
taken for the major nutrients in
high-yielding orchards and expressed per
unit of land area (Table 12.1). These esti- 12.2.1 Leaf nutrient concentration
mates indicate that the annual requirements
(for leaves and fruit) are lower than nutrient Leaf nutrient concentration reflects the fac-
requirements for many annual crops. tors influencing nutrient availability, includ-
Planting density can further alter orchard ing those affecting nutrient supply from the
nutrient demand. For example, unit-area soil, and year-to-year variation in climate
potassium requirements were greater in and crop load. Nutrient concentration is not
more densely planted ‘Golden Delicious’ stable within the season, as the rate of nutri-
orchards cited in Table 12.1 because of the ent supply and internal tree cycling alters
higher yield of fruit (which has a high potas- throughout the period of annual leaf and
sium concentration). Few nutrient estimates shoot development. Some nutrient concen-
are available for ‘super-spindle’ plantings, trations decrease (e.g. nitrogen, phosphorus,
which can have tree densities in excess of potassium and zinc), while others, such as
4000 trees ha1 and usually have even calcium and manganese, increase over the
higher fruit yields from a greater number of growing season (Fig. 12.1). As a conse-
small trees per unit land area. quence, a standard sampling time is used
when annual concentration changes are min-
imal for most nutrients. Generally this
12.2 Direct Determination Of Apple- period occurs 110–125 days post full bloom,
tree Nutrient Status which coincides with the middle of the
growing season.
In orchards, direct determination of leaf Since considerable variation in leaf nutri-
nutrient concentration has most frequently ent concentration can be measured among
been used as a routine method of monitoring various types of leaves and even leaf posi-
and adjusting the nutritional status of apple tions within a tree and among trees within
trees. Recently, with increased concerns for an orchard block, standard sampling proce-
Table 12.1. Distribution of major nutrients in apple orchards.
‘Delicious’a ‘Golden Delicious’b

(30 years old, 124 trees ha1, (14 years old, 500 trees ha1,
44.8 t fruit ha1) estimated 90 t fruit ha1)
Nutrient Whole Top Fruit Whole

kg ha1 tree framework Leaf Roots crop tree
Nitrogen 110.5 39.7 32.6 27.6 21.3 121.2

Phosphorus 17.8 6.0 3.9 5.6 4.0 19.5
Potassium 141.7 33.9 25.7 16.8 120.0 196.4
Calcium 167.7 83.5 53.5 21.0 4.4 162.4
Magnesium 25.4 8.0 6.7 3.5 3.7 21.9
Sulphur – 8.5 2.8 3.9 0.2 15.4
Chlorine – 4.5 24.3 1.3 15.2 45.3
aAs adapted from Batjer et al. (1952); whole-tree values include blossoms, fruitlets and prunings
removed from the tree.
bAs adapted from Haynes and Goh (1980); leaf measurements at leaf fall; framework and root
measurements in dormant season.

Nutritional Requirements of Apple 269
3.50 35
(all nutrients except Zn)

3.00 30
Zn (µg g–1 dry weight)

Dry weight (%)
2.50 25
2.00 20
Ca
1.50 15
Zn K
1.00 N 10
0.50 Mg 5
0.00 P
0
20 50 80 110 140 170 200 230
Days from full bloom
Fig. 12.1. Seasonal trends of nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg)
and zinc (Zn) concentration of mid-terminal, new-year extension-growth leaves for ‘Delicious’ apple (data
from Rogers et al., 1953, except for Zn data from Neilsen, 1988).
dures are usually prescribed to obtain con- Care is usually taken to select leaves free
sistent and representative samples. Samples of insect, disease and mechanical damage,
commonly comprise 25–50 leaves collected with surfaces uncontaminated by chemical
from 20–25 randomly selected trees from residues. Standard leaf values are usually
the same cultivar/rootstock combination, compiled for heavily cropping trees, since
with leaves collected around the tree from light crops, such as those resulting from
the mid-shoot portion of the current sea- biennial bearing, differentially affect leaf
son’s extension growth on shoots of repre- nutrient concentration, decreasing, for exam-
sentative vigour (Fig. 12.2). ple, leaf calcium while increasing leaf potas-
Fig. 12.2. Typical mid-terminal, new-year extension leaves suitable for mid-season sampling.
sium. Considerable practical advice concern- limited (Righetti et al., 1990) and DRIS fre-
ing the preparation and chemical analysis of quently provides no more information than
collected samples is available from publica- the use of critical values.
tions concerned with tissue testing (Jones,
1998). Critical leaf nutrient standards applic-
able for apple are available (Table 12.2). 12.2.2 Fruit nutrient concentrations
Under ideal conditions, such standards
would be developed for each apple cultivar In some fruit-growing regions, fruit mineral
and climatic condition, since there are impor- analysis, especially for calcium, is used as an
tant genetic and environmental factors aid in postharvest management decisions
affecting nutrient uptake and requirements (e.g. Perring, 1984). As with leaf values, indi-
and the expression of deficiency. vidual fruit nutrient concentrations vary
Nevertheless, these values provide general considerably. As illustrated for nitrogen and
guidelines for assessing the nutritional status calcium, there are major differences in nutri-
of apple. ent concentrations among cultivars and
A different approach to interpreting leaf years and within the season (Fig. 12.3).
nutrient concentrations is the diagnosis and Furthermore, within- and between-tree vari-
recommendation integrated system (DRIS), ation, often associated with variation in size
which is based upon comparing nutrient of individual apples, frequently necessitates
ratios between sample plants and a high- sampling large numbers of fruit (25–50) in
yielding subgroup (Beverly, 1991). This order to obtain values representative of the
method purports to be less affected by time crop. Fruit nutrient concentrations have been
of sampling and to rank nutrients in order of expressed on a fresh weight basis after wet
their limitations to growth. Experience with digestion or on a dry-weight basis after ash-
orchard crops, especially apple, has been ing freeze-dried samples. It is also important
Table 12.2. Critical leaf and fruit nutrient concentrations for apple.
Nutrient Unit Deficiencya Normal Toxicity
Leaf
Nitrogen % DW < 1.5 1.7–2.5
Phosphorus % DW < 0.13 0.15–0.30
Potassium % DW <1 1.5–2.5
Calcium % DW < 0.7 1.2–2.0
Magnesium % DW < 0.20 0.26–0.36
Sulphur % DW < 0.1 0.1–0.3
Manganese p.p.m. DW < 25 25–120 > 120
Iron p.p.m. DW < 45 (?)b 45–500
Boron p.p.m. DW < 20 20–60 > 70
Copper p.p.m. DW <5 5–12
Zinc p.p.m. DW < 14 15–120 130–160
Molybdenum p.p.m. DW < 0.05 0.1–0.2
Fruit (harvest)c
Nitrogen mg 100 g1 FW 50–70
Phosphorus mg 100 g1 FW 7–9d > 11
Calcium mg 100 g1 FW <4 >5
Boron mg 100 g1 FW < 0.8
aSeverely deficient leaves can sometimes have unexpectedly higher leaf concentrations.
bLeafiron concentrations not correlated with iron deficiency.
cWhole-fruit concentrations minus stems and seeds.
dAppropriate for cultivars susceptible to low-temperature breakdown.
DW, dry weight; FW, fresh weight.

(a)
1000
900 ‘McIntosh’
N concentration (mg kg–1) ‘Spartan’
800
700
600
500
400
300
9 6 3 0 9 6 3 0
Weeks before harvest Weeks before harvest
Year 1 Year 2
(b)
‘McIntosh’
120 ‘Spartan’
Ca concentration (mg kg–1)
100
80
60
40
20
6 3 0 9 9 6 3 0
Weeks before harvest Weeks before harvest
Year 1 Year 2
Fig. 12.3. Seasonal trend in (a) fruit nitrogen (N) and (b) fruit calcium (Ca) a fresh-weight concentration as
affected by cultivar in successive years. Average of data from 30 commercial orchards in the Pacific north-
west. Vertical bars represent mean value ±1 standard error when larger than symbol. (Adapted from Wolk et
al., 1998.)
to consider the part of the fruit that has been 12.3 Nutritional Implications of Rooting
analysed, since values are reported for whole Characteristics
fruit (usually without seeds), skin, cortical
plugs, opposite sectors and so on. Nutrient The root system plays a major role in the
gradients occur from skin to core and from absorption and translocation of water and
calyx to stem end of fruits and nutrient redis- nutrients from the soil throughout the tree.
tribution can occur postharvest. Faust et al. Apple cultivars are usually grafted on clonal
(1967) indicated patterns that differ by nutri- rootstocks, which have been selected on the
ent for apple fruit. Some critical values pro- basis of characteristics such as precocity, abil-
posed for whole apple fruits are indicated in ity to reduce scion vigour and resistance to
Table 12.2, although it is likely that such val- pests, rather than ability to take up water
ues are less universally applicable than criti- and nutrients (Chapter 5). Nevertheless,
cal leaf nutrient concentrations. apple root systems have several general
characteristics that affect their nutrition and 1–2 m and, without competition from other
response to soil conditions. trees, can achieve a lateral spread exceeding
The distribution and effectiveness of fruit that of the top branches. Despite the poten-
tree roots, including apple, have been com- tial for extending great distances and
prehensively reviewed by Atkinson (1980). depths, apple root density is low, frequently
Apple root systems have been mapped after orders of magnitude less than that of
relatively labour-intensive excavations (Fig. Graminaceae species, with which apple is
12.4). Roots are often non-uniformly distrib- often interplanted (Fig. 12.5). This implies a
uted within the exploitable soil volume, can more limited exploitation of the soil volume
sometimes penetrate to depths exceeding than might otherwise be expected.
Fig. 12.4. Excavated whole apple tree on vigorous rootstock, East Malling Research Station (photo courtesy
D. Atkinson).
Length of root per area of soil surface (cm cm–2)
10 4 10 3 10 2 10 1
herbaceous
Graminaceae
herbaceous
non-Graminaceae
woody plants
apple
Fig. 12.5. Typical range of root length, expressed as cm cm2 of soil surface, for various species, including
apple.
There is also considerable plasticity in the Shoot

growth of apple root systems. For example,
Soil
roots proliferate when nutrient and water con-
ditions are favourable, as beneath drip emit- 2
ters through which nutrients are applied. In
loamy sand soils, average root location, after 5 3
years of nitrogen and phosphorus fertigation
through drip emitters, was within 30 cm of the
soil surface and emitter location for ‘McIntosh’
Root
apple trees on M.26 and M.9 rootstocks (Fig.
1
12.6). The pattern was less pronounced for the
more vigorous M.7 rootstock. Atkinson (1980) Fig. 12.7. Major pathways of nutrient acquisition by
also indicates that planting density influences plant roots: 1, root interception; 2, mass flow of
root distribution, with roots deeper and more water; 3, diffusion (adapted from Barber, 1984).
laterally restricted when trees are planted
more closely together. 12.4 Soil Testing
Nutrient uptake by roots occurs by direct
root interception, by mass flow of dissolved Less reliance is placed upon soil testing to
nutrients in water absorbed by the plant and determine the nutritional status of apple
by diffusion if a concentration gradient for orchards. Difficulties associated with the
the specific ion develops round the absorbing meaningful use of soil tests for perennial
root (Fig. 12.7). For an annual maize crop (Zea crops such as apple are greater than for
mays L.) growing in a fertile soil, the principal annuals. It is difficult to collect a representa-
pathways have been calculated for key nutri- tive sample from the rooting zone of a crop
ents (Table 12.3) and they indicate that domi- that is both deeply and sparsely rooted. Also,
nant pathways differ for nutrients. Apple is localization of roots, as induced by fertilizer
likely to access fewer nutrients by direct and water additions, may disproportionately
interception due to lower root density and increase the importance of a small portion of
because, in general, apple trees are grown in the soil profile. Critical soil values are not
low fertility soils. Nevertheless, the relative easily established for apple, which poten-
importance of the pathways is probably per- tially has a longer period of nutrient uptake
tinent for apple and the consequences for than annuals and also has the capacity of
availability of nutrients in orchard soils is storing and recycling some nutrients within
subsequently discussed for each nutrient. a perennial root and top framework.
Distance from emitter (cm)

0 25 50
0
Depth below surface (cm)
10
20 M.9
M.26
30 M.7
40
50
Fig. 12.6. Average depth below surface and distance from a drip emitter for all roots of ‘McIntosh’ apple
as affected by rootstock. Trees maintained for five growing seasons under daily drip irrigation with annual
N + P fertigation.
Table 12.3. Principal method of nutrient adsorption for maize (adapted from Barber, 1984).
Amount of Proportion (%) of requirement suppliable by

nutrient
Adsorption required Root
method/nutrient (kg ha1) Diffusion Mass flow interception
Primarily by diffusion
Phosphorus 39 94 6 3
Potassium 196 78 20 2
Primarily by mass flow
Nitrogen 190 0 99 1
Sulphur 22 0 95 5
Copper 0.1 0 400 10
Boron 0.2 0 350 10
Molybdenum 0.01 0 200 10
Mass flow and root interception
Calcium 39 0 429 171
Magnesium 45 0 250 38
Manganese 0.3 0 133 33
Mass flow, diffusion and root
interception
Zinc 0.3 33 33 33
Most soil extracts attempt to simultane- plants, as detailed for individual nutrients
ously characterize rapidly available soluble later in this chapter. Soil salinity is important
and exchangeable nutrients while estimating because apple trees, like most fruit crops, are
the quantity of nutrients potentially available sensitive to excess salinity. Methods of deal-
from inorganic and organic soil components. ing with suboptimum pH and salinity condi-
The usefulness of these values varies with tions are discussed in Chapter 11.
their ability to simulate the soil chemical con-
ditions that control the availability of a nutri-
ent and so predict the uptake of nutrients by 12.5 Augmenting Apple Nutrition
the crop. This is likely to vary with the indi-
vidual chemistry of nutrients as discussed in 12.5.1 Soil applications
the following sections describing soil availabil-
ity for each nutrient. There have also been In apple orchards, fertilizers are applied
insufficient long-term fertilization trials for directly to the soil to raise nutrient levels if
apple to calibrate annual responses to modi- they are inadequate for the successful
fied soil nutrient levels. Important changes in growth of the crop. They are also applied to
crop load and soil moisture regimes, for exam- maintain soil fertility, which will decline if
ple, can modify responses in successive years. nutrient removal from the soil, via processes
Despite these limitations, soil testing can such as crop uptake (Table 12.1), leaching,
be a useful indicator of the relative nutrient volatilization or denitrification, exceeds
status of orchards, especially when monitor- nutrients added via weathering of minerals
ing changes over time. Furthermore, it is the and the mineralization of organic matter.
only way to determine possible nutrient lim-
itations prior to orchard establishment
12.5.1.1 Solid fertilizer additions
(Chapter 11). Several important soil charac-
teristics, including soil pH and salinity, can Nitrogen is the most frequently deficient and
be readily measured. Soil pH has an impor- most commonly applied fertilizer in
tant effect on the availability of nutrients to orchards. Addition to the soil of phosphorus
and potassium is warranted when soil-test of mono-ammonium phosphate fertilizers

results, plant response or tissue analysis within the planting hole is an example of
indicate a need. Calcium additions can be band application in orchards, which can suc-
large when lime is applied to increase soil cessfully stimulate initial tree root growth,
pH. Sometimes magnesium and boron appli- providing the salinity created by concentrat-
cations are recommended, while most other ing the soluble fertilizer is not excessive.
micronutrients are rarely applied via the soil. An important environmental effect of fer-
Additional details concerning effective soil tilization is the acidifying tendency of com-
application of various nutrients are dis- mon orchard fertilizers (Table 12.4) resulting
cussed further in the orchard-management from the conversion of ammonium to
section for each nutrient. nitrate-nitrogen in the soil.
Placement of solid fertilizers, particularly
the more insoluble forms such as phospho-
12.5.1.2 Soluble fertilizer additions
rus and potassium, is very convenient prior
(fertigation)
to orchard establishment because of the abil-
ity to readily incorporate broadcast fertilizer. The addition of fertilizers with irrigation
Soluble fertilizers, especially nitrogen, can be water (often referred to as fertigation) is a
broadcast on the orchard surface and readily newer technique for fertilizing apple trees.
carried into the root zone with precipitation Fertigation has several advantages, includ-
or irrigation. Placement, as concentrated ing the ability to transport soluble nutrients
within the herbicide strip, may be important directly to the root zone whenever water is
in order to reduce competitive uptake from applied to the plant. In this way, fertilizer
orchard-floor vegetation. Banding of essen- amounts and timing can be precise and
tial fertilizer is less frequently undertaken in adjusted to coincide more closely with actual
established orchards, despite a potential for plant demand, without the necessity of fre-
more effective uptake of such fertilizers in quent traffic throughout the orchard to
soils of low fertility. Application of high rates broadcast fertilizer. The method works best
Table 12.4. Equivalent acidity of commonly applied fertilizers and suitability for fertigation (adapted from
Tisdale et al., 1993).
Equivalent acidity
(kg CaCO3 per 100 kg
Fertilizer Content fertilizer to neutralize) Fertigated
Nitrogen
Ammonium nitrate 33–34% N 62 Yes
Ammonium sulphate 21% N, 24% S 110 Yes, acidity concerns
Urea 45–46% N 71 Yes
Calcium nitrate 15.5% N 20 Yes
Potassium nitrate 13–14% N, 26 Yes
44–46% K2O
Phosphorus
Phosphoric acid 52–54% P2O5 110 Yes, acidity concerns
Mono-ammonium phosphate 11% N, 48% P2O5 58 No
Di-ammonium phosphate 16–18% N, 70 No
46–48% P2O5
Triple super phosphate 45–46% P2O5, Neutral No
1% S
Potassium
Potassium chloride 60–62% K2O Neutral Yes
Potassium sulphate 22% K2O, 22% S Neutral No
with low-pressure micro-irrigation systems, Details regarding the principles and prac-
including drip, microjet and minisprinkler, tices of fertigation are beyond the scope of
which tend to concentrate roots in smaller this chapter but are available in several
soil volumes due to localized water addi- recent reviews (Haynes, 1985; Bar-Yosef,
tions. This, in turn, increases tree reliance on 1999). Since optimum water application is
a smaller soil volume, creating a necessity to critical to correct delivery of fertigated nutri-
maintain optimum soil conditions in this ents, much valuable information can also be
important zone. Declining pH can be a seri- obtained from standard irrigation manuals
ous problem, especially when acidifying fer- and texts, such as that prepared for drip irri-
tilizers are repeatedly applied to poorly gation by Dasberg and Bresler (1985).
buffered soils. The advantages of targeting
nutrient additions directly to the root zone
and the concentrating effects on root distrib- 12.5.2 Foliar applications
ution are not observed when fertigation
occurs via sprinkler systems, which apply Nutrients can be directly supplied to apple
water over the whole orchard floor. trees via spray application of dilute concen-
Only readily soluble fertilizers (Table trations of minerals to foliage, buds and
12.4) can be fertigated after inclusion via an even bark. The quantity of nutrients capa-
injector or siphon into the irrigation system ble of being absorbed through waxy outer
(Fig. 12.8). Fertilization can thus be adjusted
by irrigation zone but not to suit individual
trees. Comparative studies between broad-
cast and fertigated application of nitrogen
usually emphasize the ability to achieve
similar yield, growth or nitrogen uptake at
lower nitrogen rates when fertigating
(Neilsen et al., 1999). Effective scheduling of
irrigation to avoid over-application of water
(as discussed later in this chapter for nitro-
gen) offers the potential to reduce leaching
of fertigated nitrogen to the groundwater,
since nitrogen moves with water.
The mobility of phosphorus and potas-
sium is much greater when fertigated,
increasing the ability to apply these nutri-
ents rapidly when required. Experience with
fertigating micronutrients is quite limited.
Although chelates or sulphates of most
minor elements are sufficiently soluble to
fertigate, these nutrients, if required, are
most efficiently supplied via foliar sprays.
Incomplete understanding of the seasonal
variation in nutrient demand by apple cur-
rently limits growers from taking maximum
advantage of the flexibility of fertigation
timing. The attractiveness of fertigation will
increase as knowledge of tree nutrient
demand improves. Additional information
regarding the fertigation of various nutri-
ents as part of an orchard nutrient-manage-
ment strategy is discussed separately for Fig. 12.8. Fertigation headworks for addition of
individual nutrients. soluble fertilizer directly into the irrigation system.
cell layers is often small relative to tree 12.6 Special Nutrient Considerations
nutrient demand. Nevertheless, for apple,
timely application of several nutrients via 12.6.1 Organic production
sprays has improved tree growth and yield
by amelioration of deficiency symptoms (as There is increasing interest in organic apple
for micronutrients) but has also improved production, as discussed in Chapter 22. Such
fruit quality (as for calcium and phopho- an orientation can alter nutritional practices
rus) by reduction of physiological disorders since allowable fertilizer products are regu-
of fruit (Table 12.5). The mechanisms of lated by an organic certification body.
nutrient uptake and factors improving Although there can be some variation in
absorption of foliar-applied nutrients have acceptability of products among groups, in
been detailed in a review by Swietlik and general, use of synthetic fertilizers is often
Faust (1984). The relative importance of not allowed and there is a greater emphasis
foliar application as a source of supply of on maintenance of soil organic matter via
individual nutrients when managing use of organic composts, mulches and green
orchard nutrition is subsequently discussed manures (Edwards, 1998). Thus nitrogen, for
for each nutrient. example, is not applied as a fertilizer such as
Table 12.5. Timing, rate and concentration of major individual mineral nutrient sprays successfully
applied to apple.
Spray
Ratea concentration
Nutrient Common form Usual timing (kg ha1) (g 100 l1)
Major nutrients
Nitrogen Urea To foliage to correct deficiency, including 2–11
postharvest to maintain cropping 200–1000
Magnesium Epsom salts To early-season foliage (during 45–90 1200–2000
(MgSO4.7H2O), extension growth) to correct deficiency
Mg(NO3)2 22.5–45 500–600
Phosphorus KH2PO4 To early-season foliage and fruit to 22 1000
reduce low-temperature breakdown
Calcium CaCl2 To fruit, usually mid- to late season to 14–21 300–500
reduce disorders such as bitter-pit and
Ca(NO3)2 improve quality 23–34 600
Micronutrients
Boron Solubor Before pink and to foliage, including 2.8–5.6 60–100
postharvest to maintain yield and
correct deficiency
Zinc ZnSO4 (36% Late dormant (silver tip to bud swell) 4.5–45 120–1200
solid) as maintenance or to correct deficiency
ZnSO4 (liquid 18.9–100 l 250–3000 ml l1
form, 47 g Zn l1),
Zn chelates, To foliage to correct deficiency, Manufacturer’s label
Zn oxides maintain levels guidelines
Iron Fe chelates To foliage to correct deficiency 1.1–2.25 Label guidelines

Manganese MnSO4 To foliage to correct deficiency 2–9 60–200
Copper Copper To early-season foliage of non-fruiting 1–2 50
oxysulphate, trees; late dormant (green tip) in
copper fruiting trees 1–2
oxychloride 200
aHigher rates may be required when deficiency is severe and multiple applications may also be required.
ammonium nitrate but rather as an appro- 12.7 Individual Nutrients

priate organic source with lower nitrogen
content, which is slowly released after min- 12.7.1 Nitrogen
eralization of nitrogen-containing organic
compounds. Acceptable nitrogen sources Nitrogen is a highly mobile nutrient that
might include, for example, fish fertilizers, cycles between the atmosphere, the soil, liv-
blood meal or various composted organics, ing organisms and ground- and surface
providing they do not contain prohibited water (Fig. 12.9). It is the nutrient that is
constituents, such as sewage biosolids. most often applied to apple trees and in the
Similarly acceptable sources of phosphorus largest amounts and, consequently, has been
may include suitable composts and rock the most studied.
phosphate, while potassium might be
applied via composts, wood ashes or a
12.7.1.1 Soil availability
mined but not synthesized mineral, such as
potassium sulphate. Similar restrictions and Inorganic nitrogen is taken up by tree roots
recommendations exist for most mineral as either ammonium or nitrate. To a lesser
supplements, including addition of amend- extent, apple roots are also capable of
ments to adjust soil pH. Orchardists con- absorbing organic nitrogen compounds,
templating organic production should be including urea, glutamate and aspartate. The
aware of the regulations appropriate to availability of soil nitrogen for plant growth
their region and be prepared for the addi- is dependent on both organic and inorganic
tional emphasis on the long-term mainte- soil properties and on factors determining
nance of soil fertility. microbial activity. The majority of nitrogen
taken up by the plant is as nitrate (Mengel
and Kirkby, 1982), which occurs naturally in
12.6.2 Integrated fruit production soil solutions in much higher concentrations
(up to 200 p.p.m.) than ammonium (~1
Recently there has been a trend in commer- p.p.m.), and this relationship persists even
cial fruit production towards optimization when soils have been amended with ammo-
of apple growth and quality while safe- nium fertilizer (Fig. 12.10). In soils, nitrate is
guarding the environment and human unbuffered – that is, it is present almost
health. This is often referred to as integrated entirely in solution; thus the majority of
fruit production (IFP) (see Chapter 21) and nitrate moves to the tree root in mass flow,
also has implications concerning orchard although localized diffusion gradients may
nutritional practices. Some issues, such as arise when depletion zones develop around
maintenance or improvement in orchard soil roots. In contrast, ammonium is adsorbed to
fertility and organic-matter content, parallel the soil cation-exchange complex and can
the concerns of organic production. There is also be fixed within the lattices of certain 2 :1
a further desire to control nutrient and layer clay minerals, such as illite and vermi-
water-supply to minimize effects on the culite, often competing with potassium for
environment. This is likely to mean such sites. Movement of ammonium to the
increased chemical analysis of orchard soil, root is by both mass flow and diffusion.
leaf and fruit samples to target precisely The concentration of nitrate within the
nutrient application to demand. Avoiding soil solution is governed by the microbial
excessive water application will also reduce mineralization of organic matter and the
nitrate contamination of groundwater. conversion of ammonium to nitrate (nitrifi-
Fertigation has been suggested as more cation), nitrate uptake by microorganisms
appropriate to IFP, since nutrient application and plants and nitrate leaching by water
rates can be reduced and timing matched to from precipitation or irrigation (Fig. 12.9).
tree nutrient demand. It is likely that further Additions of nitrogen to the soil occur by
implications for nutrient management will microbial fixation of N2 from the atmos-
emerge as practices of IFP evolve. phere, dissolved in precipitation, or by inor-
NH3 N2
N2,
Crop N N2O, NO
removal Industrial
fixation
N2
Organic amendments
NO3–, NH4+, Fertilizer
Organic N NH3
NH4+, NO3–
Biological
fixation
Uptake of Mineralization
NO3–, NH4+ Denitrification
Organic
Organic matter
matter NH4+
Nitrification
Immobilization H+ Soil acidification
–
and degradation
NO33–
NO
Leaching
Fig. 12.9. Potential cycling of nitrogen in a typical apple orchard.
300
260
Start of End of
220 fertigation fertigation
Nitrate-N (p.p.m.)
(31 May) (9 Aug.)

180
140
100
60
20
140 160 180 200 220 240 260 280

Day of the year
Fig. 12.10. Change in soil solution nitrate-N concentration at 30 cm depth beneath drip emitters in
response to daily fertigation (31 May–9 Aug.) of 30 g N as either ammonium sulphate (—● ) or calcium nitrate
) . Vertical bars represent mean value ±1 standard error.
(—
ganic fertilizer and organic amendments. As availability of nitrogen for growth. For
a consequence, the concentration and avail- example, the nitrification of ammonium did
ability of nitrate in the soil solution vary not occur immediately on addition of
considerably throughout the growing sea- ammonium sulphate to orchard soils (Fig.
son. Factors that affect microbial activity, 12.10) but was probably suppressed by low
such as temperature, carbon source, oxygen soil temperatures and incorporation into
and water-supply, all have a bearing on the soil organic matter.
Nitrogen additions as organic amendments tion in a soil receiving spring-broadcast fertil-

or fertilizer can have several detrimental izer and sprinkler irrigation (Fig. 12.11) indi-
effects on the environment. The high mobility cated that availability of broadcast fertilizer
of nitrate results in losses to groundwater, par- may be limited by rapid leaching of applied
ticularly in well-drained soils receiving either nitrogen out of the root zone. Similar effects
high levels of precipitation or irrigation. Soil might be expected if a broadcast fertilizer
application of ammonium- or urea-based fer- application is followed by a heavy rainfall.
tilizers may result in acidification as a result of
nitrification (Table 12.4), particularly in poorly
12.7.1.2 Tree nitrogen demands
buffered soils. Gaseous losses of ammonia to
the atmosphere are most likely to occur when It has long been recognized that tree fruits
ammonium-based fertilizers are surface- have a requirement for nitrogen that may be
applied to high-pH soils, whereas denitrifica- greater than that supplied by the soil.
tion is most likely in waterlogged situations Increased vegetative growth responses of
where soils are poorly drained. apple trees were demonstrated by Lyon et al.
Because the state of soil nitrogen is so (1923) and increased bloom by Bradford
dynamic, limited use has been made of soil (1924) in proportion to additions of fertilizer
tests to determine availability. Extractions with nitrogen. Plant nitrogen is a major constituent
2 M KCl have been used to determine both of amino acids, proteins, nucleic acids and
nitrate and ammonium levels in soils. other compounds and plays a major role in
Weinbaum et al. (1992) point out that, despite plant metabolic processes. Nitrogen uptake
the difficulties associated with assessing poten- by roots is an active process and is indepen-
tial nitrogen mineralization in tree root zones dent of concentration and maximum uptake
and the inherent variability in root and soil by apples can occur at very low soil-solution
nitrate distribution, high residual nitrate levels concentrations (~0.28 p.p.b.) (Bhat, 1983).
in orchard soils should be heeded as a signal to Thus, restricted uptake at low soil-solution
reduce fertilizer inputs. Changes in soil-solu- concentrations is probably the result of an
tion nitrate and ammonium concentrations inability to maintain the movement of suffi-
over time and in response to both fertilizer and cient nitrogen to the root surface. Nitrate is
water inputs can also be monitored through reduced to ammonium mainly within the
soil suction lysimeters. Sampling of soil solu- roots in apple trees, although movement of
100
Start of End of
fertigation fertigation
80 (25 May) (29 June)
Nitrate-N (p.p.m.)
60
40
20
0
140 150 160 170 180 190 200 210 220 230 240
Day of the year
Fig. 12.11. Change in soil solution nitrate-N concentration during the growing season in response to 25 g
nitrogen as ammonium nitrate, applied either as a single broadcast application on 25 May (— ● ) or as eight
) (25 May–29 June ). Drip irrigation applied daily. Vertical bars represent
fertigations made every 5 days (—
mean value ±1 standard error.
root-supplied nitrate into leaves and subse- total tree nitrogen was found in leaf and fruit
quent reduction have been reported for apple tissue, of which 18% could be attributed to
trees receiving high levels of nitrate fertilizer fruit (Neilsen et al., 2001b). Attempts have
(Titus and Kang, 1982). been made to determine the annual demand
The absolute nitrogen requirements of for nitrogen based on the removal of nitro-
trees are high compared with other nutrients gen from the orchard. In mature trees (14–21
(Table 12.1). The total nitrogen content of years), around 25–26 kg nitrogen ha1 was
apple trees shows some variation ranging removed annually in fruit and prunings
from around 2 g per tree at planting for high- (Greenham, 1980; Haynes and Goh, 1980).
density apple plantings of trees on dwarf When the portion of nitrogen annual
rootstocks to up to 890 g per tree for stan- uptake that is incorporated into the frame-
dard 30-year-old trees (Tables 12.1 and 12.6). work of the tree was included, annual removal
In modern (high-density) production sys- was estimated as 33 kg ha1 from 9–12-year-
tems, total tree nitrogen content by year 6 is old ‘Cox’s Orange Pippin’/M.7 (Greenham,
probably around 30 g per tree (100 kg ha1). 1980) and 52 kg ha1 from 30-year-old
On an annual basis, nitrogen is required ‘Delicious’ trees (Batjer et al., 1952). These esti-
to support the growth of new tissues. The mates assume that all fallen leaf tissue is incor-
developing leaves and fruit of apple trees are porated back into the soil. However, in
major sinks for nitrogen. In mid-season, it semiarid areas where orchard floors may be
has been estimated that 40% of total tree dry, fallen leaves may be blown out of tree
nitrogen may be found in leaf tissue in stan- rows and thus may not be incorporated back
dard mature apple trees (Batjer et al., 1952) into the tree root zone. Under this assumption
and 40–50% in leaves of young potted trees nitrogen removal may then include losses
(Forshey, 1963). In young dwarf trees, 51% of from senescent leaves (Table 12.7).
Table 12.6. Total nitrogen content of apple trees.
Nitrogen content
Tree density
(trees ha1) Cultivar g per tree kg ha1
3300 ‘Golden Delicious’/M.9 at planting 2.2 7.4

3300 ‘Golden Delicious’/M.9 end of year 1a 8.2 27
3300 ‘Elstar’/M.9 end of year 4b 19.7 65
n/a M.9/M.9 end of year 5c 15.0 n/a
aNeilsen et al. (2001a).
bNeilsen et al. (2001b).
cMason and Whitfield (1960).
n/a, not available.
Table 12.7. Estimates of annual nitrogen requirements of dwarf apple

trees: nitrogen removal in fruit and senescent leaves.
Nitrogen content
Cultivar g per tree kg ha1a
‘Golden Delicious’/M.9 end of year 1 2.84 9.4

‘Elstar’/M.9 end of year 4 10.21 33.1
‘Gala’/M.9 end of year 3 10.49 34.7
‘Gala’/M.9 end of year 6 12.19 40.2
aAssumes tree density of 3330 trees ha1 in the autumn and that tree
rows characteristically have weed-free strips.
Estimates of nitrogen requirements for cence, but it is likely that the most rapid
growth are complicated by plant nitrogen withdrawal occurs 3–4 weeks before leaf fall.
cycling, which occurs both within and The timing and rate of nitrogen withdrawal
between seasons. Extensive reviews of this probably depend on many factors, including
topic have been previously published (Titus nutrient availability, crop load and environ-
and Kang, 1982; Millard, 1996). Nitrogen mental conditions. Nitrogen withdrawn
assimilated by leaves is stored throughout from leaves in the autumn is stored in the
much of the growing season as leaf protein, woody tissues of the tree as proteins or
but, as with most deciduous trees, some of amino acids, which are broken down and are
the stored nitrogen can be mobilized and subsequently remobilized to supply nitrogen
withdrawn from apple leaves before leaf for the growth of new tissues the following
abscission. The proportion of leaf nitrogen spring. The total amount of tree nitrogen that
that is withdrawn from apple leaves during is involved in remobilization may be quite
senescence has been measured in several large. In newly planted ‘Golden
studies and ranges from 23 to 50% (Titus and Delicious’/M.9, over 50% of total tree nitro-
Kang, 1982). The withdrawal of leaf nitrogen gen at planting had been cycled out of
may start as early as the cessation of shoot woody tissues to support new root and shoot
growth or as late as the onset of leaf senes- growth 70 days later (Fig. 12.12a).
(a) 1400
1200 Woody tissue

N content (mg per tree)
1000
800
600
Planting date
400 April 9th
200 Shoots
Roots
0
90 100 110 120 130 140 150 160 170 180 190 200
Day of the year

(b) 4
N content (mg per leaf)
3 Root-supplied N
Remobilized N
1
0
110 120 130 140 150 160 170 180
Day of the year
Fig. 12.12. (a) Remobilization of unlabelled nitrogen from woody tissue into new growth in newly planted
‘Golden Delicious’/M.9 (dotted lines represent period when unlabelled nitrogen could be from both
remobilization and root uptake). (b) Content of root-supplied and remobilized nitrogen (N) in spur leaves of
3-year-old ‘Elstar’/M.9. Vertical bars represent mean value ±1 standard error.
Early spring remobilization of nitrogen is Undersupply of nitrogen leads to poor

largely independent of current nitrogen sup- tree growth and reduced yield due to fewer
ply to the roots (Tromp and Ovaa, 1973; and smaller fruit. Symptoms of nitrogen
Millard, 1996). In general, evidence suggests deficiency include poor fruit set and early
that root uptake of nitrogen in the spring fruit drop, small leaves that are uniformly
does not occur until after remobilization is pale green to yellow, chlorotic basal leaves
under way. For 3-year-old ‘Elstar’/M.9 trees, that abscise prematurely and thin twigs with
the nitrogen content of spur leaves came brown to reddish bark. Diagnosis of tree
mainly from remobilization and was present nitrogen status is often based on measure-
in the leaves before root uptake of nitrogen ments of shoot growth and leaf nitrogen con-
occurred (Fig. 12.12b). This may be because centration in midsummer, with subsequent
low soil temperatures in the spring reduce nitrogen requirement based on a reference
nitrate uptake or because high concentrations range of concentrations where < 1.5% is con-
of remobilized and cycling amino acids in the sidered deficient and 1.7–2.5% is considered
roots and/or a lack of necessary carbon sufficient (Table 12.2).
skeletons for amino acid synthesis suppress In general, oversupply of nitrogen is more
nitrogen uptake. Thus remobilized nitrogen of a problem in orchard management than
is probably most important to the growth undersupply. In addition to potential leakage
that occurs before the development of the to the environment of excess nitrogen that is
shoot-leaf canopy with its substantial photo- not taken up by trees, oversupply of nitrogen
synthetic capability. The relative importance can lead to excessive vigour and poor fruit-
of remobilized nitrogen to seasonal growth is quality characteristics (Faust, 1989). Typical
dependent on the magnitude of both the effects may be reduced firmness at harvest
nitrogen in storage and that which is cur- and after storage or reduced red colour
rently available. In the absence of root-sup- development when internal ethylene pro-
plied nitrogen, a strong relationship has been duction continues to occur, resulting in fruit
shown between the extent of shoot growth that are picked over-mature (Fallahi, 1997).
and the amount of stored nitrogen (Harley et The incidence of several storage disorders,
al., 1958; Hill-Cottingham and Bollard, 1965). such as cork spot, bitter pit and internal
In M.26 apple rootstocks the proportion of browning and breakdown (Bramlage et al.,
nitrogen for leaf growth that was derived 1980; Terblanche et al., 1980), has also been
from remobilization varied between 18 and linked to excess nitrogen supply. Such detri-
87% for trees receiving high and low supplies mental effects of excess nitrogen have been
of nitrogen in the spring, respectively linked to increases in fruit size and shading
(Millard and Neilsen, 1989). Remobilized from vigorously growing shoots, among
nitrogen contributed 50% of the nitrogen in other factors. Vigorous growth related to
shoot leaves, 90% of the nitrogen in the spur nitrogen oversupply has also been linked to
leaves that subtend the fruit and 60% of the increased susceptibility to diseases, such as
nitrogen in the fruit (Neilsen et al., 2001b). fire blight (Van der Zwet and Keil, 1979).
The time when root nitrogen uptake starts to
occur is thus probably a function of tree
12.7.1.3 Orchard nitrogen management
nitrogen status (demand), phenological stage
and soil nitrogen availability. Later in the sea- Extensive reviews of orchard nitrogen man-
son, the timing of nitrogen supply to the agement and its effect on the environment
roots can affect partitioning of nitrogen to dif- have been published previously (Weinbaum
ferent tissues. Khemira et al. (1998) showed et al., 1992; Sanchez et al., 1995). Both of these
that spring-applied broadcast fertilizer in 9- reviews recommend that fertilization rates
year-old standard and spur-type and management strategies for tree fruits
‘Delicious’/M.7 trees was found in above- should include a judgement of tree nitrogen
ground tissues rather than roots, but that status, crop nitrogen demand, soil nitrogen
nitrogen from a preharvest broadcast applica- availability and other site-specific variables.
tion was allocated preferentially to the roots. Responses to nitrogen fertilizer additions,
such as increased vigour, increased yield, remobilization (Fig. 12.12) and nitrate can be
darker leaf colour and higher leaf nitrogen rapidly removed from the root zone when
concentration, may only occur with trees of broadcast fertilizer applications are followed
low nitrogen status, as nitrogen uptake is by irrigation (Fig. 12.11) or by heavy rainfall.
probably restricted in trees of high nitrogen Autumn broadcast applications are also
status (Table 12.8). Moreover, leaf nitrogen likely to be inefficient, although this would
concentration may be unaffected by depend on timing with respect to root activ-
increased uptake of nitrogen in response to ity. Applications made in colder regions,
supply if dilution due to increased leaf where uptake may be limited by tempera-
growth occurs. Thus, leaf nitrogen concen- ture and demand, are likely to be susceptible
tration as a sole method of diagnosis of nitro- to leaching by either winter rains or snow
gen requirement is not satisfactory, melt (Tagliavini et al., 1996). The effects of
particularly when tree nitrogen status is nitrogen on apple cold-hardiness are unclear.
above deficiency. Estimates of tree nitrogen Concern is often expressed that late-season
demand based on removal in the crop and applications of nitrogen, which may promote
other tissues, as previously described, could new extension growth, could result in
be used as a starting-point for fertilizer increased susceptibility to winter damage.
application rates, modified by consideration Conversely, trees with low nitrogen status
of leaf and soil nitrogen concentration. may also be sufficiently weakened to be sus-
Compared with many other crops, the ceptible to winter damage. However, there
annual nitrogen requirement for apple trees are few published data to either support or
is a relatively modest 30–100 kg ha1. For refute these statements. Because of the inher-
established trees, there are also numerous ent inefficiency in single-dose nitrogen appli-
reports of adequate growth and cropping cations, typical application rates of up to 200
without any nitrogen-fertilizer additions on kg ha1 may be four times the tree require-
fertile soils (Atkinson, 1980). However, rec- ments (Sanchez et al., 1995). More efficient
ommended nitrogen application rates may use of nitrogen is likely if broadcast applica-
typically range from 75 to 150 kg ha1 and in tions are split to match periods of demand.
some cases even higher. Traditionally, grow- Khemira et al. (1998) demonstrated that late-
ers have broadcast nitrogen fertilizer in summer broadcast nitrogen applications
spring and/or autumn. Single-broadcast (preharvest) did not adversely affect current
applications of nitrogen imply that either fruit nitrogen concentrations but resulted in
sufficient nitrogen can be taken up during a root uptake and storage of nitrogen, which
restricted period of high nitrogen availabil- was subsequently remobilized for growth in
ity, just after application, or that applied the following season. The amount of nitro-
nitrogen becomes incorporated into soil gen fertilizer required is also affected by
organic matter and is slowly released. competition from orchard-floor vegetation.
However, single-dose, spring fertilizer appli- Decreases in the amount of broadcast nitro-
cations may be inefficient, as uptake in gen required to meet growth requirements
spring can be limited by low demand during have been reported in response to the intro-
Table 12.8. Estimates of fertilizer nitrogen applied in excess of crop nitrogen demand (adapted from
Klein et al., 1989, and Weinbaum et al., 1992).
Nitrogen Leaf nitrogen Mean annual Estimated annual Estimated mean

application rate concentration yield removal in crop nitrogen excess
(kg ha1) (%) (t ha1) (kg ha1) (kg ha1)
50 1.9–2.4 47.7 23.8 26.2

150 2.4–3.2 54.3 27.1 122.8
250 2.5–3.1 45.1 23.0 226.9
400 2.5–3.2 47.1 23.5 376.4
duction of weed-free (herbicide or mulch) much closer than typical broadcast rates to
strips associated with the tree row (Hogue estimates of tree demand (Table 12.7) for
and Neilsen, 1987). high-density apple orchards.
An alternative method of fertilizer supply Control of nitrogen losses through leach-
is through irrigation water (fertigation), ing may be more achievable in irrigated than
which has been used both where irrigation is in rain-fed orchards, particularly if micro-irri-
mandatory and where it is supplemental. gation systems are used and water applica-
Usually applied through low-pressure tions are scheduled to meet evaporative
microsystems, fertigation of nitrogen offers demand. Under sprinkler irrigation, leaching
the possibility of controlling availability both losses of up to 242 kg ha1 have been
spatially and temporally and potentially reported for apple trees receiving up to
improving nitrogen-fertilizer use efficiency. 175 cm rainfall + irrigation per year
Controlled availability over time is evident (Stevenson and Neilsen, 1990). Leaching
from measurements of soil-solution nitrate losses from trees receiving daily drip irriga-
concentrations in response to daily (Fig. tion were much lower when irrigation was
12.10) and weekly fertilizer applications (Fig. scheduled to meet evaporative demand
12.11). Levin et al. (1980) suggested that it is rather than applied at a fixed rate (Fig. 12.13).
appropriate to assume a 50% nitrogen use In order to control the timing of nitrogen
efficiency for fertigated apple trees. supply and potentially reduce total inputs
Comparisons of fertigated and broadcast and losses to the environment, a consider-
applications of nitrogen on tree growth and able amount of work has been undertaken
yield show, in general, that lower rates of fer- on supplying nitrogen by foliar sprays (Titus
tigated nitrogen maintain a similar tree nitro- and Kang, 1982). Urea is the most common
gen status to that achieved with higher rates nitrogen form in foliar nutrition (Table 12.5),
of broadcast nitrogen (Neilsen et al., 1999). although calcium and potassium nitrate
Quite low rates of fertigated nitrogen were have also been successfully used. Generally,
sufficient to meet tree requirements in The solution concentrations of urea are 5% or
Netherlands (15–20 g per tree) (Kipp, 1992) lower. Translocation of summer-applied
and the UK (20 g per tree; 26 kg ha1) foliar urea from leaves to other tissues has
(Hipps, 1992) and, in both studies, tree per- been variously described as rapid and rela-
formance was either similar to or better than tively complete (Boynton, 1954) to limited
that of trees receiving broadcast nitrogen at (Forshey, 1963). Attempts to supply total tree
rates of 80–100 g per tree. The rates applied nitrogen by foliar applications have resulted
to the fertigated trees in these studies are in poor growth, possibly because of poor
8.0
Scheduled
6.0 Unscheduled
N loss (g per tree)
N
4.0
2.0
0
May June July Aug. Sept. Oct. Oct.–May
Fig. 12.13. Seasonal nitrogen (N) losses beneath the root zone for 2-year-old ‘Gala’/M.9 apple trees in
response to irrigation either applied to meet evaporative demand (scheduled) or at a fixed rate
(unscheduled). Vertical bars represent mean value ±1 standard error.
translocation out of the leaves (Forshey, are very low, often in the p.p.b. range. This is
1963). Autumn application of foliar urea a consequence of the propensity for phos-
makes use of the natural mechanism of with- phorus absorption and precipitation in the
drawal of nitrogen from leaves during senes- soil, especially as calcium phosphates at soil
cence to augment tree nitrogen status for pH above 7 and as iron and aluminium
growth the following spring. Increased ovule phosphates below pH 4. Optimum phospho-
longevity the following spring has been rus availability thus generally occurs in the
demonstrated with autumn applications of mid-pH range between acid and basic pH
foliar urea. In pot studies with young trees, conditions. Soils also vary, however, in their
foliar nitrogen withdrawal ranged from 23 to ability to maintain adequate phosphorus in
70% (Titus and Kang, 1982). However, soil solution (a soil’s phosphate buffer capac-
Khemira et al. (1998) reported that, in mature ity (PBC)). High-PBC soils require much
spur and standard ‘Delicious’ trees, very lit- higher phosphorus additions than low-PBC
tle nitrogen derived from autumn foliar soils to achieve the same soil-solution phos-
applications was found in any tissues the fol- phorus concentration. High-PBC soils also
lowing season. There is some evidence that, require higher rates of phosphorus fertilizer
for trees with sufficient nitrogen at senes- in order to achieve effective downward
cence, foliar-applied urea may only replace movement of phosphorus into the main root-
leaf nitrogen that would normally be with- ing zone in orchards.
drawn, rather than augmenting it. Thus As a result of low soil-solution phospho-
autumn applications of urea would be most rus concentration and the low rooting den-
useful in trees with low nitrogen status. sity of apple trees, phosphorus uptake in
In summary, supplying fertilizer nitrogen apple is strongly dependent upon desorption
to meet tree requirements should take into of phosphorus from the soil matrix and its
account the management of nitrogen derived diffusion to tree roots. Consequently, soil
for growth both from plant storage and from properties such as low temperature and
root uptake. The natural cycling of nitrogen in moisture contents, which decrease desorp-
the tree described above gives the opportu- tion and diffusion, also decrease phosphorus
nity to manipulate tree nitrogen status in both uptake. It has been difficult to develop a
the current and the future growing season. meaningful soil extract that adequately sim-
Fertigation offers a relatively efficient way to ulates all factors affecting phosphorus
supply nitrogen if water is well managed. For uptake for apple, whose roots are also
trees receiving broadcast nitrogen, a low- known to be infected by vesicular-arbuscular
input strategy for nitrogen management may endomycorrhizae, which in themselves stim-
be to reserve ground applications in the ulate tree phosphorus uptake. As a result,
limited use has been made of various
spring for trees with very low nitrogen status
extractable soil phosphorus determinations
and to supply nitrogen either on the ground
to design phosphorus-fertilization pro-
after harvest and/or by foliar urea spray in
grammes. Orchards with soils low in
the autumn to trees of adequate nitrogen sta-
extractable phosphorus, regardless of the
tus. In all cases, the need for nitrogen fertilizer
extractant used, are most likely to respond to
applications should be based on tree perfor-
phosphorus fertilization. In contrast, apple
mance and, where possible, measurement of
growth has been stimulated via application
residual nitrate concentration in the soil. of high rates of phosphorus in the planting
hole at sites otherwise testing at medium
and even high phosphorus levels.
12.7.2 Phosphorus
12.7.2.1 Soil availability 12.7.2.2 Tree phosphorus demands

Phosphorus solubility is low in most soils, The absolute phosphorus requirements of
with the result that concentrations of readily apple trees are small relative to other nutri-
available soluble phosphorus in soil solution ents (Table 12.1). Plant phosphorus is a factor
in energy transfer and is a constituent of deficient trees, adequate mid-terminal-leaf

phytic acid storage compounds and nucleic phosphorus concentrations are between 0.20
acids. Limited research on phosphorus and 0.30% dry weight (DW), usually exceed-
uptake characteristics of apple rootstocks ing thresholds developed for mature fruiting
under field conditions by Bhat (1983) indi- trees (Table 12.2). Desirable fruit phosphorus
cated that solution phosphorus concentra- concentrations are likely to vary by cultivar
tions at which phosphorus inflow ceases to but have also been suggested for cultivars
increase are higher for apple than for most (Table 12.2) where an association between
other species. Thus, maintenance of high low fruit phosphorus and reduced fruit qual-
soil-solution phosphorus for extended peri- ity has been established.
ods of time should result in high phosphorus
uptake. High phosphorus demand in apple 12.7.2.3 Orchard phosphorus management
trees occurs during periods of considerable
meristematic activity (of which phosphorus The traditional belief has been that apple is
is an important regulator) as roots and unlikely to respond to phosphorus fertiliza-
shoots emerge, particularly at planting. tion, as documented in numerous phospho-
Hansen (1980) has indicated from pot experi- rus-fertilizer trials (Childers, 1966).
ments using young ‘Golden Delicious’/M.9 Nevertheless, sufficient responsive sites have
that total phosphorus uptake is about 50% now been identified from a range of fruit-
lower in fruiting than in non-fruiting trees. growing areas for the possibility of growth
Nevertheless, annual phosphorus require- and yield responses to surface application of
ments can be high early in the growing sea- superphosphate or mono- and di-ammo-
son in fruiting trees when extensive cell nium phosphate not to be ruled out for
division is occurring in developing leaves apples grown on soils testing very low in
and fruitlets at a time when root length is at extractable phosphorus.
a minimum. Increased incidence of low-tem- Application of high rates of soluble ammo-
perature breakdown of stored fruit has been nium-phosphate fertilizers directly in the
associated with low phosphorus concentra- planting hole has been widely used in the
tions of certain cultivars, including ‘Bramley’ Pacific Northwest of North America as a
and ‘Cox’s Orange Pippin’ apples in the UK method of improving the establishment and
and ‘McIntosh’ apples in North America. accelerating the vegetative growth and initial
Unequivocal indicators of insufficient yield of newly planted apple trees. The
phosphorus have rarely been observed in the method seems particularly effective where
field on apple. Taylor and Goubran (1975), soils are being replanted and have previously
working with phosphorus-deficient apple been fumigated. Phosphorus is usually
trees in Australia, indicated that symptoms applied at high rates (100–150 g per planting
are frequently an expression of reduced rates hole), well dispersed in the soil to avoid salt
of meristematic activity and include accumu- toxicity, and can be effective over a range of
lation of soluble nitrogen and anthocyanins, extractable soil phosphorus. Similar initial
resulting in dark green leaves with purple growth and yield improvements have been
tints in autumn. Also noteworthy was achieved by fertigation of relatively high rates
retarded bud burst and reduced flowering of of phosphorus (10–20 g P per tree) early in the
phosphorus-deficient trees. Similarly, Benson establishment year. Both of these soil-applica-
and Covey (1979), working with ‘Golden tion methods result in temporary elevation of
Delicious’ apple growing in nutrient solution, soil-solution phosphorus concentration in the
indicated that phosphorus deficiency was main rooting zone and take advantage of the
characterized by reduced leaf size and shoot previously described potential for high phos-
growth but showed no visual, morphological phorus uptake of apple roots.
signs of deficiency. These observations indi- Foliar application of soluble phosphorus
cate that it can be difficult to diagnose phos- compounds has successfully augmented the
phorus deficiency from field symptoms. phosphorus nutrition of apple. Sprays have
From studies involving young phosphorus- usually been applied to improve fruit qual-
ity early in the season (within 4–6 weeks of 12.7.3.2 Tree potassium demands
bloom) during the period of fruit cell divi-
sion. For example, multiple weekly sprays Whole-tree K demands of apple are similar
of 1% potassium dihydrogen phosphate in magnitude to those of nitrogen (Table
augmented fruit phosphorus concentration 12.1), with leaf concentrations second to
and reduced low-temperature breakdown in nitrogen and fruit concentrations exceeding
certain cultivars, including ‘Bramley’, ‘Cox’s all other mineral nutrients. Potassium is
Orange Pippin’ and ‘McIntosh’ (Table 12.5). mobile within the phloem, so that fleshy
fruit are well supplied with potassium.
Fruit are such strong sinks for potassium
12.7.3 Potassium that whole-tree partitioning studies indi-
cate that fruiting trees have higher potas-
12.7.3.1 Soil availability sium uptake per unit of root dry mass than
non-fruiting trees. At the same time, heavy
The total potassium content of most soils crop loads decrease leaf potassium concen-
can be as high as 4% (average 2%) but soil- tration.
solution concentrations can be quite low, Severe potassium deficiency results in
since much of the potassium is unavailable symptoms typified by marginal leaf
to plants due to its incorporation within the browning and scorch on older basal leaves of
structure of many soil minerals. Soluble new year extension growth, the mobile
potassium is in equilibrium with potassium nutrient potassium being preferentially
adsorbed on negatively charged exchange translocated to younger leaves (Plate 12.1).
sites on the surfaces of clay minerals and Symptoms appear from mid- to late summer
organic matter. Soils with high contents of as fruit potassium demands increase. At
certain clay minerals, including illite and advanced deficiency, leaves can have
vermiculite, have a particular affinity for irregular edges as dead tissue breaks off.
strongly adsorbing potassium within their Spur leaves subtending fruit can also be
structure. Such soils with ‘high potassium- affected, developing an irregular chloro-
fixation capacity’ require higher rates of tic surface during midsummer, which
potassium application to achieve the same progresses into interveinal browning and
increases in soil-solution potassium concen- marginal leaf scorch by fruit harvest. Since
tration as could be achieved by application potassium is the major positively charged ion
of lower rates to soils with low clay content. absorbed by apple, compensatory uptake of
In an analogous manner to phosphorus, other positively charged ions (especially
potassium supply to plant roots depends magnesium and calcium) occurs under
mainly on the diffusive flux that a soil can conditions of severe deficiency. Potassium-
maintain in the direction of plants. Thus, deficient trees are therefore often
light-textured soils with high sand contents characterized by unusually high leaf mag-
and a tendency to droughty conditions are nesium concentrations. Severe leaf symptoms
more likely to be potassium deficient, espe- commonly occur when leaf potassium
cially for apple trees, which have low root concentrations are less than 1%. Such
densities. The potassium status of orchard reductions in leaf photosynthetic capability
soils can be approximated by determination frequently reduce fruit size and yield.
of the potassium concentration in a variety Amelioration of potassium deficiency can
of soil extracts, including neutral, 1 M increase fruit red colour and titratable acidity,
ammonium acetate (Grimme and Nemeth, but such effects are often not apparent when
1978). Soils that result in potassium defi- tree potassium status is adequate. Potassium
ciency of apple usually have low extractable deficiency can also impair tree water
potassium. However, as with many soil relationships, because of the role potassium
tests, ‘available potassium’ is often poorly plays in the proper functioning of stomata.
correlated with apple tree potassium uptake Frequently, apples with calcium-related
over a range of soils and climates. physiological disorders, such as bitter pit or
poor storage quality, have high fruit potas- 12.7.4 Magnesium

sium/calcium ratios. However, calcium con-
tent appears to be the critical factor and the 12.7.4.1 Soil availability
effect on storage and fruit quality of
Magnesium, like potassium, occurs in the
increased fruit potassium concentration
soil in different forms, including the rela-
when calcium supply is adequate is unclear.
tively unavailable magnesium contained in
Elimination of potassium deficiency, how-
the structure of soil minerals, exchangeable
ever, improves fruit yield and quality.
magnesium adsorbed on organic matter and
clay minerals and soluble magnesium dis-
solved in the soil solution. Unlike potassium,
12.7.3.3 Orchard potassium management magnesium is not susceptible to specific fixa-
It is generally recognized that moderate to tion within the structure of clay minerals.
heavy broadcast application of potassium The proportion of magnesium as a fraction
salts to the soil surface increases avail- of all exchangeable cations is usually second
able potassium to the rooting depth in to calcium and exceeds that of potassium. As
most orchard soils (Boynton and Oberly, a consequence, soil-solution concentrations
1966). Most potassium salts appear to be are relatively high and mass flow, rather
suitable potassium sources, including potas- than diffusion, is capable of supplying the
sium chloride, potassium sulphate, potas- magnesium needs of apple trees. Magnesium
sium–magnesium sulphate and potassium availability is, however, affected by the ionic
nitrate, with some advantages to the applica- balance between it and other positively
tion of sulphate salts when high rates are charged cations, including K+, Ca2+, NH4+,
required or soils are saline. Details of the Mn2+ and H+. Thus magnesium deficiencies
positive responses of apple to soil-applied may be induced in heavily limed or potas-
potassium, often at rates of 50–100 kg potassium fertilized soils. Soil pH also affects
sium ha1, have been reviewed (Forshey, magnesium availability, which is usually
1969; Cummings, 1985). Less certain are ben- optimum in slightly acid soils of pH 5–7.
efits from potassium application when leaf Below pH 5, H+ and Al3+ antagonisms are
concentrations exceed the low range (above common, while, above 7, Ca2+ predominates.
1.5%). However, it is probably important to The availability of magnesium varies with
moderate potassium applications in order to soils, usually being lower for coarse-textured,
ensure a favourable calcium/potassium bal- leached soils formed on parent materials con-
ance, which will optimize quality and stora- taining few magnesium-containing minerals
bility of the apple fruit. Of the leaf nutrients, and high on soils where leaching is low and
magnesium is most adversely affected by high-magnesium parent materials, such as
high applications of potassium. basalts or dolomites, predominate. It is possi-
Fertigated potassium is readily available in ble to estimate plant-available magnesium by
most soils, including those which normally using an extractant, such as neutral, 1 M
‘fix’ large quantities of broadcast potassium ammonium acetate, to measure exchangeable
(Uriu et al., 1980). For example, annual fertiga- and soluble magnesium. In this way, soil-test-
tion of 15 g K per tree successfully prevented ing laboratories can distinguish orchard soils
the development of growth-inhibiting potas- low in available magnesium and susceptible
sium deficiency in drip-irrigated high-density to the development of magnesium deficiency.
apple orchards on sandy soils (Neilsen et al.,
1998).
12.7.4.2 Tree magnesium demands
Few instances have been observed where
apple trees have responded to foliar potas- The apple tree’s quantitative requirements
sium applications – in part, because of the for magnesium are less than for potassium
large quantities of potassium required for and calcium (Table 12.1). An important
optimum growth and the success in amelio- fraction of total plant magnesium (usually
rating deficiency via soil application. 15–20%) is required as a constituent of the
chlorophyll molecule and the ion serves example, application of magnesium-contain-

important biochemical functions in activat- ing dolomite limestone is likely to be effec-
ing enzymes involving phosphorylation, tive in acid soils with low exchange capacity
activation of ribulose bisphosphate car- and less effective where soil pH and calcium
boxylase and protein synthesis (Mengel content are high. Similarly, applications of
and Kirkby, 1982). magnesium sulphate (250 kg Mg ha1) did
Deficiencies of magnesium are most not eliminate symptoms for deficient trees in
apparent in leaf tissue. Symptom expression the short term when soil potassium levels
varies with the severity of the disorder and were high (Woodbridge, 1955) or for defi-
the susceptibility of the apple cultivar. For cient rootstocks when low annual rates
example, ‘McIntosh’ apple is more prone to (40–60 kg Mg ha1) were made for 8 years
magnesium deficiency than ‘Delicious’. (Ford, 1964). A caution to the indiscriminate
Interveinal leaf chlorosis and yellowing usu- application of soil magnesium in orchards is
ally develop on older leaves at the base of the increased incidence of bitter pit observed
the new year’s growth late in the season as in susceptible cultivars after the addition of
fruit size increases (Plate 12.2). Since magne- magnesium to poorly buffered sandy soils
sium is mobile in the phloem, it is preferen- (Van der Boon et al., 1966).
tially translocated to young shoot leaves and There have been few reports of effective
developing fruit. Depending on the severity fertigation of magnesium on apples, despite
of the deficiency, brown interveinal necrotic the ready solubility of various magnesium
blotches, known as leaf scorch, can develop, salts applied as foliar sprays. Multiple sprays
coalesce and, for some cultivars (e.g. (usually two to five) of magnesium salts
‘Newtown’) develop into chlorotic and have ameliorated magnesium-deficiency
‘scorched’ edges difficult to distinguish from symptoms. Such sprays are best applied
potassium deficiency. Severe deficiency can post-bloom and are usually recommended
result in leaf curling and premature defolia- for the period of rapid shoot growth (late
tion. The resulting reduction in canopy pho- May–early July in northern latitudes). They
tosynthesis can also reduce fruit size and have eliminated decreased fruit set, normally
lead to premature fruit drop. Mild magne- associated with severe magnesium defi-
sium deficiencies do not necessarily result in ciency (Ford et al., 1965). A 2% (w/v) magne-
a growth and yield depression, since the sium sulphate (Epsom salt) solution has been
photosynthetic activity of the remaining commonly used, although other soluble
unaffected green leaves can be enhanced magnesium salts, including magnesium
(Ford, 1966). In general, magnesium defi- chloride and magnesium nitrate, have
ciency is unlikely when leaf magnesium con-
worked (Table 12.5). Some soluble magne-
centrations exceed 0.26%, while leaves with
sium chelates have been ineffective because
concentrations below 0.20% are likely to
of low magnesium concentration (Neilsen
exhibit the magnesium deficiencies previ-
and Hoyt, 1984).
ously described (Table 12.2).
12.7.4.3 Orchard magnesium management 12.7.5 Calcium

Application of magnesium-containing mate-
rials to orchard soils can improve the long-
term magnesium nutrition of apple trees, as Most soils contain large quantities of calcium
long as a meaningful alteration in soil mag- (3.5%, 35 t ha1) as a constituent of calcium
nesium status, especially relative to the other carbonate, silicate, sulphate and phosphate
soil cations, can be achieved in the main minerals. Calcareous soils can be particularly
rooting zone (Mason, 1964). Failure to enriched in calcium, with contents as high as
achieve this may explain reported ambigui- 10–20%. Calcium usually comprises the bulk
ties concerning the effectiveness of soil mag- of exchangeable cations (65–85%) adsorbed
nesium applications (Boynton, 1947). For to organic matter and inorganic soil colloids
and has the highest concentration (50–100 mal skin bronzing, late-season darkening of
p.p.m.) of any cation in soil solution. Thus, lenticels and sometimes severe fruit splitting
plant requirements for calcium are usually by harvest. It is now well established that
satisfied by mass flow of water to the root, inadequate calcium supply is related to
and low soil calcium is not often a limitation many physiological disorders in the storage
to growth. An exception can occur for old, organs of fruits and vegetables (Shear, 1975).
leached and acidic soils, which may have For apples, calcium-related disorders include
extremely low calcium contents. The pres- bitter pit (Plate 12.3), cork spot, cracking,
ence of high concentrations of other cations internal breakdown (Plate 12.4), Jonathon
also inhibits uptake of calcium. Soil calcium spot, lenticel blotch, water-core and low-tem-
status is intimately associated with soil pH, perature and senescent breakdown. In gen-
since higher calcium saturation of the eral, improved calcium nutrition is also
exchange complex occurs as pH increases. considered to provide broad-spectrum pro-
Soils can be tested for their extractable cal- tection against many postharvest pathogens.
cium content, but pH measurements usually It is therefore not surprising that consid-
determine the necessity of applying calcium- erable effort has gone into understanding the
rich liming materials to the soil. inflow of calcium into apple fruit. Different
seasonal patterns have been measured, rang-
ing from the bulk of inflow occurring in the
12.7.5.2 Tree calcium demands
4–6-week period of cell division following
Apple wood contains more calcium than any bloom (Wilkinson, 1968) to a steady increase
other mineral element, with the result that throughout the growing season (Faust, 1989).
orchard requirements to maintain top and Regardless of the temporal pattern of
root structures are higher than for all other absolute calcium accumulation, calcium con-
nutrients (Table 12.1). Calcium transport to centration declines as fruit size reaches a
the root surface within the soil is usually maximum at harvest. Concentration gradi-
high but, within the plant, calcium move- ents also develop within the fruit, with mini-
ment is slowed by ion exchange in the mum values measurable immediately
xylem. Calcium is also relatively immobile in beneath the skin at the calyx end of the fruit.
the phloem, due to restricted phloem load- Actual calcium concentrations achieved in a
ing. As a consequence, plant organs more given year are therefore strongly affected by
dependent upon phloem supply, such as the final fruit size and hence crop load. A large
fruit of the apple, frequently have difficulties yield tends to produce smaller fruit of higher
obtaining sufficient amounts. Calcium serves calcium concentration. A strong genetic
important functions within the plant, includ- effect is also apparent, as different cultivars
ing regulation of cellular behaviour and can achieve different fruit calcium concentra-
maintenance of cell integrity and membrane tions despite growing under similar environ-
permeability (Mengel and Kirkby, 1982). mental conditions. This is illustrated for
In apple, the effects of inadequate levels different cultivars growing in the irrigated
are manifested primarily in fruit rather than fruit-growing region of the Pacific north-
leaves. Leaf deficiency symptoms are rarely west of North America (Fig. 12.3).
seen under orchard conditions but have been The rarity of leaf calcium deficiency in the
described for apples grown in nutrient solu- orchard and the frequency and importance of
tions containing low calcium concentrations. fruit calcium disorders indicate that fruit cal-
The descriptions of Shear (1971) for the culti- cium concentration thresholds are a more
var ‘York Imperial’ are pertinent and include useful indicator of an orchard’s calcium sta-
first an upward cupping of the youngest tus. Unfortunately, such values are likely to
shoot leaves, followed by the development be cultivar-specific and dependent upon the
of veinal and interveinal chlorosis, with the analysis method and the part of the apple
eventual development of chlorotic spots and analysed. For example, recommended whole-
necrotic tissue on leaf edges. Fruit symptoms fruit calcium concentrations (Table 12.2) may
may be more prominent and include abnor- not be pertinent for ‘Cox’s Orange Pippin’,
which has a deficiency threshold of 5 mg Ca unimportant. The amount of calcium

100 g1 fresh weight. A critical peel calcium absorbed by the fruit is directly related to the
concentration of 700 mg kg1 DW was rec- number of sprays applied, so the number of
ommended for ‘Baldwin’ apple, although recommended sprays depends upon the cal-
year-to-year variation in critical calcium val- cium status of the fruit. Four or five sprays at
ues was also apparent (Drake et al., 1974). 7–10-day intervals late in the growing season
have eliminated serious occurrences of bitter
pit in susceptible cultivars growing in semi-
12.7.5.3 Orchard calcium management arid climates where little precipitation occurs
Considerable effort has been made to under- to wash the applied calcium from the fruit.
stand factors influencing orchard calcium Late-season sprays are particularly effective,
nutrition, because of the importance of this since the intended fruit target is larger and
nutrient for fruit quality. Many of the details since many calcium disorders develop late in
have been reported in comprehensive the season or postharvest during storage.
reviews on the subject (Vang-Petersen, 1980; Early-season sprays may be appropriate for
Himelrick and McDuffie, 1983). calcium-related disorders, such as corking,
Applying large quantities of calcium com- which develop early in the season.
pounds to the soil via liming can increase Calcium can also be applied postharvest
leaf calcium concentration but not fruit cal- to picked fruit by dipping, vacuum or pres-
cium concentration, as indicated by persis- sure infiltration with CaCl2 salts with con-
tence of calcium-related physiological centrations as high as 2% (w/v). The
disorders despite the liming of acidic soils. advantages of such treatments to reduce
Thus, lime applications are made primarily postharvest decay have been described in
for pH adjustment, rather than as a source of detail by Conway et al. (1992).
calcium. Soil application of soluble calcium
salts, such as calcium nitrate, is unlikely to
alter the calcium status of most fruit, except 12.7.6 Sulphur
in very acid soils (Van Lune, 1984). Similarly,
successful use of fertigation of soluble cal- 12.7.6.1 Soil availability
cium to increase fruit calcium concentration Most sulphur contained in soils (90–95%)
has not been reported, possibly because the occurs in organic compounds, while the
calcium concentration of fertigating solu- remaining inorganic forms usually occur as
tions is often less than the calcium concentra- sulphates in normally aerated orchard soils.
tions measured in soil solution. Inorganic forms are readily leached in humid
Successful augmentation of fruit calcium regions and thus can occur in high concentra-
concentration and diminution of calcium- tions in ground and runoff waters. There is lit-
related disorders have been achieved by tle effect of soil pH on sulphur availability. No
application of multiple sprays of soluble cal- single extraction technique has found univer-
cium salts directly to the fruit surface prior to sal acceptance for analysing available sulphur
harvest. Calcium chloride is usually recom- in soils for most crops (Kowalenko, 1993),
mended, although Ca(NO3)2 can also be effec- with the consequence that little information is
tive (Table 12.5). The chloride form is cheap, available to relate the performance of apple
contains a higher calcium concentration and trees to the supply of soil sulphur. Inadequate
avoids additional application of nitrogen to availability of sulphur is most likely to occur
fruit. Some materials containing, for example, in orchards with leached soils, low in organic-
chelated calcium at low concentrations have matter content and receiving limited sulphur.
been ineffective as foliar calcium sources.
Recommended concentrations of the salt are
12.7.6.2 Tree sulphur demands
usually about 0.5% (w/v) but single-spray
applications as high as 4% have been effec- Apple requirements for sulphur are similar
tively applied just prior to harvest when the in magnitude to those for phosphorus, with
resulting leaf damage has been assumed to be absorbed SO4-sulphur incorporated struc-
turally into sulphur-containing amino acids, ments also have significant sulphur content,
proteins and coenzymes. Thus deficiencies of approximating 10% of their nitrogen content.
sulphur inhibit protein synthesis and reduce
tree growth. Sulphur deficiencies have rarely
been observed on apples in the field. A 12.7.7 Boron
notable exception was described by Benson
et al. (1963) in Washington State. Deficient Boron has been the subject of numerous
apple trees exhibited a chlorosis on younger review articles, including a recently pub-
leaves at the top of the plant, with leaf colour lished book (Gupta, 1993), which provides
progressing from light green, light yellow to additional useful information concerning
yellow, eventually affecting the whole plant, this nutrient.
stunting growth and reducing yield. The
trees did not respond to nitrogen or iron 12.7.7.1 Soil availability
applications, deficiencies of which can result
in similar symptoms. Severe sulphur- Total boron contents of orchard soils nor-
deficiency symptoms on shoot-tip leaves mally range from 20 to 200 p.p.m., often with
developed when tissue SO4-sulphur concen- only a small portion (< 5%) of the total avail-
trations were less than 100 p.p.m. (0.01%). As able to plants. Boron, dissolved in soil solu-
a result of the rarity of sulphur deficiency, tion as the uncharged boric acid molecule, is
there is some question as to how this concen- plant-available but is also easily leached
tration relates to the more easily measured from sandy soils. Organic matter can contain
total sulphur value, although a critical total appreciable quantities of boron, which are
sulphur concentration of 0.1% has been available to plants upon mineralization.
suggested (Table 12.2). Boron adsorption to clay and organic matter
increases with pH, reaching a maximum
sorption and minimum solution concentra-
12.7.6.3 Orchard sulphur management tion at pH 8–9. Thus leached sandy soils that
Many orchards have substantial natural inputs are low in organic matter are susceptible to
of sulphur with precipitation and irrigation. boron deficiency, particularly after liming. In
Unintended applications of sulphur can also arid regions, high concentrations of soluble
occur via sulphur-containing fertilizers and boron salts can accumulate in soil or irriga-
pesticides. For example, sulphate salts of tion waters, creating a potential for toxicity.
ammonium (24% S), calcium (19% S), potas- The amount of boron extracted after boiling
sium (17–22% S), magnesium (14% S) and zinc soil in water is commonly used as a measure
(11% S), as well as superphosphate fertilizer of plant availability. Values below 1 p.p.m. or
(12% S), may be applied as a source of other above 5 p.p.m. indicate potential for defi-
nutrients but simultaneously augment sul- ciencies or toxicities of boron, respectively.
phur. Some concern has been expressed that
sulphur inputs to orchards are decreasing as 12.7.7.2 Tree boron demands
control of air pollution reduces the sulphur
content of precipitation and the use of sul- In apple, boron is required at low (p.p.m.)
phur-containing fertilizers and pesticides in concentrations in leaves and fruit. In general,
orchards declines. boron maintains plant meristematic activity
Amelioration of inadequate sulphur and cell-wall stability and functions as a
nutrition was demonstrated in deficient coenzyme in the formation and transport of
Washington State orchards via both broad- sucrose. Apple has less sensitivity to boron
cast application of sulphur-containing fertil- deficiency than other tree fruits, such as
izers to the soil and spray applications of peach. However, deficiency of boron in apple
soluble sulphate salts to foliage. It is also has serious consequences, which are mani-
likely that soluble sulphate salts could be fested first in reproductive structures, includ-
effectively fertigated, although little research ing flowers and fruit. Inadequate boron
has been reported. Most organic amend- nutrition has been associated with ‘blossom
blast’, a drying and shrivelling of flowers at health at leaf boron concentrations as low as
bloom, distinguishable from frost damage by 50 p.p.m. (Hansen, 1981) but more generally
the longer retention time of the damaged tis- in excess of 70 p.p.m.
sue on the tree (Plate 12.5). Since pollen-tube
growth is also stimulated by boron, reduced
12.7.7.3 Orchard boron management
fruit set under conditions of low boron status
can further contribute to yield reductions. An overriding consideration in the manage-
Insufficient boron has also been associated ment of boron nutrition in orchards is the nar-
with fruit disorders. Commencing soon after row range between boron deficiency and
fruit set, water-soaked areas develop and toxicity. Boron can be effectively applied to
progress into dry, hard, corky masses within orchard soils to correct deficiency as various
the fruit, which can result in external depres- soluble boron fertilizers, including borax
sions and corking and cracking of the (sodium tetraborate, 11% B). Rates are rather
extended skin surface (Plate 12.6). These dis- low, usually only 1–2 kg B ha1 year1.
orders can develop early on fruit less than Application of rates as low as these can be
half final size and may result in early drop of physically difficult, so sometimes higher rates
affected fruits, further reducing yield. This are applied once every 3 years or applications
disorder has also been referred to as drought are made as boronated forms of other
spot, since boron deficiency can be induced macronutrient fertilizers. Uniform application
by low-moisture conditions preceding a is important to avoid toxic concentrations. The
period more favourable to tree growth. With addition of organic amendments and com-
increased severity of the deficiency, abnormal posts containing boron can also provide suffi-
growth of shoot tips occurs and the youngest cient boron for the soil to prevent deficiency.
leaves are misshapen and wrinkled and Given the solubility of many boron com-
develop red veins and interveinal chlorosis. pounds, the nutrient can also be fertigated in
Death of the terminal bud and small areas of irrigation water, although care must be taken
the inner bark and cambium near the grow- to avoid over application. Rates as low as 0.3
ing tip results in the dieback of shoots. It is g B per tree, applied over several weeks, have
also likely that root tips suffer dieback, but effectively ameliorated boron deficiency with-
this often goes unnoticed. Severe deficiency out inducing toxicity. It is prudent, however,
can result in rough and cracked bark (apple in irrigated areas to know the boron concen-
measles) due to the death of some bark tissue
tration of irrigation waters and therefore the
in young branches.
amount of boron already applied with water.
Boron is the nutrient with the narrowest
Foliar sprays of soluble boron compounds,
of margins between deficiency and toxicity.
such as Solubor (20% B), are highly effective
Apple is moderately sensitive to boron toxic-
and are often safer than soil applications,
ity, which is manifested by delays in flower-
ing and increased incidence of bud death since much lower rates and concentrations of
(blind buds). Midrib yellowing of leaves compounds (0.2–0.5% w/v) can be applied.
(Plate 12.7), defoliation, shoot dieback and Solubor is also compatible with other com-
early maturity, increased respiration and monly applied oils, emulsions and pesticides.
reduced storage life of fruits are characteris- A common strategy for maintenance of boron
tics of toxicity in apple. in apple orchards is to make a single spray
There is likely to be some genetic varia- early in the spring, prior to bloom (Table
tion in the sensitivity of different apple culti- 12.5). This can be useful, particularly if polli-
vars to deficit and toxic concentrations of nation conditions are suboptimal. Recently,
boron. Nevertheless, boron deficiency has sprays applied postharvest in the autumn
frequently been reported at leaf boron con- have also been effective the following spring
centrations < 20 p.p.m. and fruit-flesh boron as a consequence of the mobility and storage
concentrations < 0.8 p.p.m. (fresh weight) of sprayed boron in apple trees (Brown and
(Table 12.2). Toxicity thresholds are less cer- Shelp, 1997). Several sprays may be required
tain, but concern has been expressed for bud to correct severe boron deficiency.
12.7.8 Zinc Columbia orchards, found soil zinc extracted

by 0.25 M MgCl2 related more closely to zinc-
A comprehensive review of soil, plant and deficiency symptoms of apple than the more
management factors associated with zinc widely used diethylenetriaminepenta-acetic
nutrition of horticultural crops, including acid (DTPA) extract (Table 12.9).
apples, has recently been completed by
Swietlik (1999).
12.7.8.2 Tree zinc demands
Apple has a low requirement for zinc but is
also highly susceptible to zinc deficiency.
Total soil zinc contents of orchard soils are Zinc functions in several plant enzyme sys-
very low, ranging from 10 to 300 p.p.m., and tems and several plant biochemical func-
average about 80 p.p.m. Total zinc contents tions, including pH regulation in plant cells,
relate to parent material, with soils originat- protection from O 2 damage and synthesis of
ing from basic igneous rocks usually higher RNA and tryptophan – a precursor of
in zinc than soils derived from silica-rich indoleacetic acid, which is involved in shoot
minerals. Zinc occurs in the soil as relatively elongation. Symptoms of zinc deficiency fre-
insoluble forms within the structure of pri- quently occur in early spring and include
mary and secondary minerals or precipitated chlorosis (yellowing) of the youngest shoot
on solid surfaces of carbonates and iron and leaves, which are often somewhat under-
manganese oxides. sized (Plate 12.8) and narrower than normal
Forms more available to plants include (a condition referred to as ‘little leaf ’).
exchangeable zinc, associated with reactive Mobility of zinc within the plant is poor, so
clay and organic matter, or zinc dissolved in little zinc is translocated from old to young
the soil solution. Zinc, like phosphorus, leaves. Yellowing occurs between leaf veins
occurs at very low (p.p.b.) concentrations in and is less symmetric than similar symptoms
solution, so that zinc supply to the root observed on basal leaves with manganese
depends upon diffusion, rather than mass deficiency. Zinc deficiency may also result in
flow, and is adversely affected by factors blind bud and rosetting (small basal leaves
inhibiting diffusion. Soil pH strongly affects forming on the shortened terminals and lat-
zinc availability, decreasing zinc solubility eral shoots of the current season’s growth).
100-fold per unit increase in pH. Thus zinc Extreme deficiency, which is rare, can result
deficiency is commonly associated with neu- in shoot defoliation and dieback and the pro-
tral to high-pH soils, especially those contain- duction of undersized, misshapen and
ing free calcium carbonate. However, acid poorly coloured apples, which ripen early
and organic soils low in zinc and soils high in and lack flavour. Zinc-deficiency symptoms
phosphorus can also be zinc-deficient. can be very sporadic in occurrence, with all
Limited research has been undertaken to or only some shoots on a tree affected and
determine the best soil extraction method for with an occasional or many trees affected
predicting zinc status of apple trees, although within an orchard block (Plate 12.8). There
Neilsen et al. (1988), working with British can be considerable overlap in leaf zinc con-
Table 12.9. DTPA-extractable zinc, copper, manganese and iron

values determined for deficient, marginal and sufficient soils for non-
orchard crops (adapted from Tisdale et al., 1993, p. 318).
Zinc Copper Manganese Iron

Soil status (p.p.m.) (p.p.m.) (p.p.m.) (p.p.m.)
Probably deficient 0–0.5 < 0.2 < 1.0 0–2.5

Marginal 0.6–1.0 – – 2.6–4.5
Sufficient–high > 1.0 > 0.2 > 1.0 > 4.5
centrations of healthy and deficient trees, of the soil as a constituent of primary silicate,
although concentrations much below 14 oxide, hydroxide and carbonate minerals
p.p.m. are a cause for concern (Table 12.2). and secondary clay minerals, such as illite.
Zinc toxicity has rarely been reported for Iron has, however, such low solubility that
apple but has been observed when insufficient amounts can be supplied to plant
unsprayed leaf zinc concentrations range roots by mass flow. Iron movement to roots
from 130 to 160 p.p.m. (Orphanos, 1982). in soils is, however, increased by its ten-
dency to form more soluble complexes,
including chelates, with organic compounds.
12.7.8.3 Orchard zinc management Soil pH is an important control on iron solu-
Inadequate zinc is widely considered the bility, with Fe3+, the predominant soluble
most common micronutrient deficiency of iron form in aerated orchard soils, decreas-
apples. Despite the successful amelioration ing 1000-fold in solubility for each unit
of zinc deficiency in many agronomic crops increase in pH and reaching minimum solu-
via the application of 4–10 kg Zn ha1 as bility at pH 6.5–8.0. Iron availability is thus
zinc sulphate, limited emphasis has been extremely low in the approximately 25% of
placed on soil zinc application in apple the world’s arable land that is calcareous
orchards. Increased zinc uptake of apple has, with pH near 8.2. High bicarbonate concen-
however, occasionally been reported in trations in soil solution or irrigation water
orchards where soluble zinc applications can further reduce the availability of iron to
have been banded at much higher rates or plants. Since compacted or saturated soils
co-applied with organic amendments under can increase soil-solution bicarbonate con-
acidifying conditions. Fertigation of soluble centration, these soil conditions also increase
zinc compounds is possible but recom- the probability of iron deficiency on soils of
mended rates and techniques on a range of high pH. Uptake of iron from soil solution is
soils have not been widely developed. also inhibited by competition from high con-
In contrast, spray applications of zinc are centrations of ions such as Zn2+, Mn2+, Cu2+,
the most common practice for successful, Ca2+, Mg2+ and K+. It is for this reason that
immediate correction of zinc-deficiency high concentrations of other metals in
symptoms and are often applied as annual unbuffered sandy soils can induce iron defi-
maintenance sprays (Table 12.5). Unfor- ciency. Limited use of soil extractants, such
tunately, there is little residual effect from as DTPA, has been made to determine the
such sprays, due to the limited movement of iron status of orchard soils (Table 12.9).
zinc from sprayed to unsprayed leaves.
Various late-dormant sprays, including zinc 12.7.9.2 Tree iron demands
sulphate, have been successfully applied
but, when fruit are on the tree, chelated Iron is a constituent of the haem complex (a
forms are usually preferred, to avoid fruit naturally occurring plant chelate involved in
damage. Otherwise, there has often been lit- electron transfer in a number of important
tle practical difference in the effectiveness of plant enzymes) (Mengel and Kirkby, 1982). In
green leaves there is a good correlation
various forms of similar solubility and zinc
between iron supply and chlorophyll content.
concentration. Direct trunk injection of zinc
Inadequate iron nutrition results in abnormal
may be possible but has not been widely
chlorophyll development, so that deficiency
practised.
begins as an interveinal chlorosis on younger
leaves resulting in prominently green veins
(Plate 12.9). These leaves may become com-
12.7.9 Iron pletely white and devoid of chlorophyll and
may even scorch late in the summer. The
12.7.9.1 Soil availability resultant reduction in photosynthetic capabil-
Total iron content of soils is high, since iron, ity also reduces the weight and area of
on average, comprises about 5% by weight affected leaves, with the yield and quality of
apples adversely affected as the extent and of FeSO4 or various iron chelates (Table 12.5).
severity of deficiency increases over the Multiple sprays to actively growing shoots
whole tree. Further descriptions and causes are frequently required and can be moder-
of iron deficiency have frequently been ately effective, but must be reapplied annu-
reviewed, recently by Korcak (1987) for horti- ally. Direct injection of 1% solutions of iron
cultural crops, including apple. A wide range citrate or sulphate into the trunks of fruit
of total iron concentration has been measured trees, including apple, has corrected iron
in normal leaves (Table 12.2), but deficient deficiency for several years without serious
leaves can have a higher iron concentration damage to trunks.
than normal leaves. In contrast, ‘active’ iron
concentration (extracted in 0.5–1.0 M HCl)
can distinguish between normal and iron- 12.7.10 Manganese
deficient leaves but is not a reliable diagnos-
tic tool. As a consequence of disruption in 12.7.10.1 Soil availability
normal photosynthesis, iron-deficient leaves
often accumulate other minerals, such as Total manganese contents of orchard soils are
phosphorus, potassium and NO3 -nitrogen, usually higher than those of zinc, ranging
or organic components, such as citric acid. from 200 to 3000 p.p.m. and averaging about
Apples are less susceptible to iron deficiency 600 p.p.m. Manganese can be a constituent of
than other temperate fruit trees, including several primary and secondary minerals and
peaches and pears, and as a result there has occurs as insoluble and oxidized Mn3+ and
been little research to identify iron-efficient Mn4+ forms associated with iron in oxide and
rootstocks, as has been done for peaches. Iron hydroxide precipitates in the soil. The reduced
deficiency can be confounded by the occur- form of manganese (Mn2+) is readily available
rence of simultaneous multiple micronutrient to plants and can occur in soil solution at
deficiencies, including manganese and zinc, p.p.m. values, usually exceeding concentra-
so that iron deficiency has also been defined tions of the other important micronutrients,
as regreening after the foliar application of an zinc and copper. Plant-available manganese is
iron spray. affected by pH in a similar fashion to zinc,
decreasing in solubility 100-fold per unit rise
12.7.9.3 Orchard iron management in pH. Thus manganese is very soluble when
soil pH is below 5.5 and has low solubility in
It can be difficult to ameliorate iron defi- calcareous or limed soils with pH above 7.0.
ciency in apple via application of soluble Manganese solubility increases in moist, satu-
salts to the soil, due to their tendency to rated soils of acid pH, whereas reductions in
become unavailable to the plant via precipi- availability can occur as a result of microbial
tation. Limited success has been achieved, oxidation. Organic matter can increase or
however, by co-application of FeSO4 with decrease manganese availability, depending
organic matter and other strongly acidifying upon the stability of the compounds formed
fertilizers. Chelated iron compounds can be by their interaction. Limited use has been
more soluble in soils, although their effec- made of manganese soil tests for orchards, in
tiveness varies and their cost is high. part because of the complexity of characteriz-
Iron-EDDHA (ethylenediaminedi-O-hydro- ing a nutritional range extending from defi-
xyphenylacetic acid) is particularly effective ciency to toxicity (Neilsen et al., 1990). The
over a range of soil pH. Little research has DTPA micronutrient test is useful for deter-
been conducted on fertigating iron, although mining manganese status near deficiency in
it is likely that the general principles of effec- neutral and calcareous soils (Table 12.9).
tive soil application will prevail (i.e. use of
chelated iron or other soluble iron sources
12.7.10.2 Tree manganese demands
under acidifying conditions).
Iron deficiency can be corrected by direct Absolute manganese requirements are small,
application to the foliage of dilute solutions slightly exceeding tree demand for zinc. Like
Mg2+, Mn2+ is required in enzyme reactions 12.7.11 Copper

involving photosynthesis and carbon assimi-
lation and is preferentially translocated to 12.7.11.1 Soil availability
meristematic tissue in growing shoot tips.
Of the micronutrients required by apple,
Chloroplasts are most sensitive to man-
copper, which is contained in primary and
ganese deficiency. Consequently, manganese
secondary minerals, often has the lowest
deficiency in apple first appears as irregu-
total concentration in soil, averaging about
larly shaped, light green spots on the mar-
10 p.p.m., with values usually ranging from
gins and between the veins of basal shoot
1 to 50 p.p.m. Much higher concentrations
leaves (Plate 12.10). Interveinal loss of green
can occur in soils after additions of high-
colour frequently results in prominent green
copper materials, including municipal
veins. Only rarely does severe deficiency
biosolids, pig and poultry manures, mine
decrease leaf size and shoot growth.
wastes and copper-containing pesticides.
Furthermore, the deficiency may be tempo-
The availability of copper to plants, as with
rary, appearing in the spring, when roots are
other trace minerals, markedly decreases as
cold, and disappearing as soils warm. Leaf
pH rises above 7.0. At high pH, copper is
manganese concentration below 25 p.p.m.
strongly adsorbed to clays, iron and alu-
may indicate manganese deficiency. In con-
minium oxides and organic matter. Most of
trast, tree manganese uptake can be very
the low concentrations of copper dissolved
high when soil pH is acid, resulting in man-
in solution are organically complexed.
ganese toxicity. Internal bark necrosis, also
known as ‘bark measles’, is a trunk disorder There has been little use of soil testing to
associated with excess manganese, which determine the copper status of orchard
results in the blistering, cracking and peeling soils. The DTPA extractant has successfully
of the bark on the trunk, branches and shoots identified low and deficient copper sites for
of affected trees (Fig. 11.6). Certain apple cul- other crops (Table 12.9).
tivars, including ‘Delicious’ (especially
‘Starkrimson’), ‘Tydeman’ and ‘Fuji’, are 12.7.11.2 Tree copper demands
known to be more susceptible to this disor-
der, which commonly occurs when soil pH is Apple tree copper requirements are among
below 5.5 and leaf manganese concentration the lowest of all nutrients. Copper is, how-
exceeds 120 mg kg1. ever, required for chlorophyll synthesis and
in several copper-containing enzymes
involved in the reduction of molecular oxy-
12.7.10.3 Orchard manganese management gen. Copper deficiency of apple is rare but
Soil applications to correct manganese defi- has been reported from orchards in
ciency are generally not used for apple and Australia, England, South Africa and the
can be ineffective if the soil condition (e.g. USA. It is characterized by the development
high pH) that reduces the soil manganese of brown and necrotic areas on terminal
availability is not altered. Fertigation of solu- leaves, accompanied by upward curling and
ble manganese compounds is possible, but distortion of the leaves. The disorder is often
frequently leaf manganese concentration can referred to as ‘wither tip’ or ‘summer
be readily increased by fertigation of acidic dieback’, since initial leaf symptoms often
fertilizers (Table 12.4). proceed to defoliation and dieback of shoot
Foliar sprays of MnSO4 and chelated tips in the late summer. Severe deficiency
manganese solutions, applied directly to results in the cracking and shedding of bark
affected leaf tissue, effectively regreen and profuse sucker development from the
leaves, although multiple sprays may be tree base. Copper deficiency occurs in leaves
required if the deficiency is severe (Table with concentrations less than 5 p.p.m.
12.5). Direct trunk injection of MnSO4 will Copper toxicity has not been documented
also work; however, the preferred treatment for apple but has been observed for other
is foliar application. crops, where it has reduced growth.
12.7.11.3 Orchard copper management Copper chelates can also be used as foliar
fertilizers. Many orchards already receive
When copper-deficiency symptoms occur in foliar copper applications via copper-
apple orchards, they have been eliminated containing fungicides. Soil copper appli-
by foliar applications of dilute copper sul- cations, broadcast or via fertigation, might be
phate or copper chloride salts (Table 12.5). effective but are generally not used.
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13 Orchard-floor Management Systems
Ian A. Merwin
Department of Horticulture, Cornell University, Ithaca, New York, USA
13.1 General Principles and Objectives in Orchard-floor Management 303

13.2 Contemporary and Traditional Roles for Orchard Ground Covers 307
13.3 Practical Options for OFM 309
13.4 Ground-cover Vegetation Species, Types and Characteristics 311
13.5 Weed-control Methods and Equipment 313
13.5.1 Herbicide applications 313
13.5.2 Between-row cultivators 314
13.5.3 Within-row cultivators 314
13.5.4 Mowing equipment 315
13.5.5 Flame or steam weed suppression 315
13.6 Epilogue 315
13.1 General Principles and Objectives Commercial fruit growing requires sub-
in Orchard-floor Management stantial inputs of resources and expertise,
and crop values can approach US$100,000
The methods used to manage weed competi- ha1. It costs about US$20,000 ha1 to estab-
tion, maintain soil fertility, provide essential lish a high-density apple orchard and bring
soil nutrients and water for trees and it into production, and the profitability of
promote beneficial biological processes in fruit growing is affected substantially by
orchards influence the productivity and sus- OFM systems (White and DeMarree, 1992;
tainability of fruit plantings. The tactics and Geldart, 1994; Derr, 2001). Weed competition
strategies for managing surface vegetation for essential resources can stunt the growth
and soil resources in fruit plantings are of young trees, increase winter injury
referred to as orchard-floor management and disease susceptibility and reduce the
(OFM) systems (Hogue and Neilsen, 1987). quantity and quality of yields in mature trees
This chapter describes the general principles (Hogue and Neilsen, 1987; Welker and
and objectives for OFM, compares different Glenn, 1988, 1989; Glenn and Welker, 1989;
weed- and soil-management systems used by Merwin and Stiles, 1994). Apple trees do not
fruit growers and suggests opportunities for compete effectively with many weeds for
improving the sustainability of fruit growing soil nutrients and water, because the trees
by optimizing and integrating OFM systems. have relatively sparse root systems that do

304 I.A. Merwin
not exploit as large a portion of the available nutrient retention, improving fruit quality (see
root zone as most herbaceous weeds Plate 13.1) and maintaining or improving soil
(Atkinson, 1980). Surface vegetation beneath organic matter and structure (Haynes, 1981;
trees must usually be managed to curtail Skroch and Shribbs, 1986; Welker and Glenn,
competition for limiting soil resources. 1988; Moore et al., 1989; Johnson and
Weed competition with fruit trees can be Samuelson, 1990; Stevenson and Neilsen, 1990;
eliminated by residual herbicide applications Merwin et al., 1996). For example, after 5 years
that eradicate year-round all surface vegeta- under different OFM systems in a New York
tion. Such ‘weed-free’ OFM systems increase orchard, soil organic matter decreased sub-
the short-term availability of water and nutri- stantially in mechanical tillage and bare-soil
ents, but may be detrimental to both soil and residual pre-emergence herbicide treatments
water quality (Merwin et al., 1996; Elmore et and increased under mowed sod grass and
al., 1997; Glover et al., 1999). Long-term conser- hay–straw mulch treatments (Fig. 13.1). The
vation of soil fertility is especially important in fundamental challenge for sustainable OFM
a perennial crop system where good fruit- systems is to achieve a favourable balance
growing sites are limited and may be between the beneficial aspects of ground-
replanted continuously for decades or cen- cover vegetation or residues and the negative
turies. Stringent climatic and edaphic require- impacts of excessive weed competition.
ments of fruit trees often restrict orchards to To avoid unnecessary or wasteful
upland sites on well-drained soils near major weed-control measures, it is important to
lakes and rivers, where soil erosion and leach- distinguish between weeds (defined as
ing or runoff of agrochemicals into water undesirable surface vegetation that reduces
resources are potentially serious problems crop yield or quality by competing for essen-
(Logan et al., 1987; Weinbaum et al., 1992; tial resources without providing compen-
NRC, 1993). From the environmental and eco- satory benefits) and ground covers (defined
nomic perspectives, orchard ground covers as naturally occurring weeds, synthetic or
can provide important benefits – preventing biomass mulches, cover crops or turf that is
soil erosion, improving water infiltration and managed as a useful part of the crop system).
7
c
Soil OM (% dry weight)
6 bc
Initial OM ab
content ab
5 ab
a
2
TILLED PRE-HBC POST-HBC CRNVCH GRSOD STRMCH
Ground-cover systems
Fig. 13.1. Organic matter (OM) content (% dry-weight basis) in upper 20 cm topsoil after 5 years under
different ground-cover management systems. Treatment abbreviations: TILLED, monthly rotavating during
growing season; PRE-HBC, pre-emergence applications of paraquat, norflurazon and diuron herbicides in
April each year; POST-HBC, two post-emergence applications of glyphosate herbicide in May and July each
year; CRNVCH, ‘living mulch’ of crown vetch; GRSOD, mowed sod grass of red fescue and perennial
ryegrass; STRMCH, 10-cm-deep hay–straw mulch applied each May. Treatment means beneath different
letters were significantly different (LSD for P < 0.05). (Data are from Merwin et al., 1994.)
Orchard-floor Management Systems 305
It follows from these definitions that ground- (Gut et al., 1996; Merwin and Ray, 1997; Al-
cover vegetation may function as weeds in Hinai and Roper, 1999). Glenn and Welker
some circumstances and as useful cover (1996) showed that yield efficiency of mature
crops at other times. The essential objectives peach (Prunus persica Batsch) trees could be
in OFM are to maintain sufficient ground increased and pruning costs decreased by
cover to protect soil and water resources and manipulating sod competition and proximity
to suppress weed competition within critical to trees. Other research has demonstrated
time spans and surface areas of the orchard. that fruit quality (blush coloration, soluble
In other words, the optimal management solids, firmness and storage potential) can be
strategy – as with pest insects or pathogens – improved by moderate nitrogen and/or
is to control weeds only if and when they water deficits induced by ground-cover
surpass a predetermined threshold where competition at certain times of the growing
potential crop damage exceeds the cost of season (Hogue and Neilsen, 1987; Hipps et
control measures and the value of compen- al., 1990; Perret et al., 1994; Hornig and
satory benefits that ground cover provides Bunemann, 1995; Marsh et al., 1996; Neilsen
for the agroecosystem. These weed-manage- et al., 1999).
ment trade-offs and thresholds are not well Effective OFM requires understanding the
understood at present for most crop systems, relationship between nutrient uptake and
including orchards. root-growth periodicity in fruit trees and
The level of pest infestation where antici- competing weeds, but surprisingly little is
pated crop damage justifies the costs of con- known about this topic (Flores et al., 1998).
trol measures is known as the damage action Pioneering research by Head (1966) in non-
threshold or economic injury level for that replicated root-observation chambers indi-
pest (Pedigo, 1989). This concept is funda- cated bimodal peaks of apple root growth
mental to integrated pest management (IPM) early and late in the growing season of
and utilized widely in disease and insect south-east England. Recent research utilizing
control – but relatively neglected in weed mini-rhizotrons and soil respiration cham-
control (Coble and Morteusen, 1992). Recent bers for replicated observations of apple
studies indicate that temporal (critical times rootstock growth and respiration rates
for weed control during the growing season) revealed substantially different trends in root
and spatial (critical weed-free areas beneath growth, presumably due to climate or soil
trees) damage thresholds can be determined differences between England and the north-
for fruit trees. One study compared tree eastern USA (Psarras et al., 2000). For field-
growth, nutrient uptake and cumulative grown ‘Mutsu’ apple trees on M.9 rootstocks
yield of apple during 8 years in 2.5 m wide in a silt–loam soil, there was negligible root
strips where weeds were either eradicated growth early and late during two growing
year-round with residual pre-emergence seasons in New York, and the main phases of
herbicide applications or suppressed only root, canopy and fruit growth all coincided
from May to August with post-emergence from late May to mid-July (Fig. 13.2). These
herbicides that allowed substantial regrowth observations may help to explain why weed
of ground-cover vegetation from September control during early summer months has
to April (Merwin and Stiles, 1994). Despite been most effective in most studies, because
potential weed competition during the dor- this appears to be the period of greatest com-
mant season, there was no comparative petition between trees and weeds for soil
advantage in the completely weed-free OFM resources. Further observations of root-
system relative to the partially weed-free growth and nutrient-uptake timing with
system. In other studies, with tart cherry different rootstocks, soil types and ground
(Prunus cerasus L.) and apple, early-summer cover species are needed to refine OFM sys-
(May–July) weed suppression resulted in tems, based upon a thorough physiological
substantially better tree growth and yields and ecological understanding of tree/root/
compared with late-summer weed control weed competition for water and nutrients.
306 I.A. Merwin
Root emergence and Fruit and shoot growth

1
Fruit growth rate
rates (cm day–1)

0.8
Shoot growth rate
0.6
0.4
0.2
0
0.8
turnover (roots day–1)
New roots
0.6 Root turnover
0.4
0.2
0
10 25
Soil temperature (°C)

Soil/root respiration
(µmol CO2 m–2 s–1)
8 20
6 15
4 10
2 Soil/root respiration 5
Temperature
0 0
24/5 21/6 19/7 16/8 13/9 11/10 8/11
10/5 7/6 5/7 2/8 30/8 27/9 25/10
Day/month
Fig. 13.2. Relative timing and rates of shoot and fruit growth, root growth and turnover, soil/root respiration
rates and soil temperature beneath 4-year-old apple trees on M.9 rootstocks, under field conditions in a
herbicide strip during one growing season (data from Psarras et al., 2000).
Water and nitrogen are the essential Keppel, 1985; Wijsmuller and Baart, 1988;
resources most often implicated in competi- Merwin and Ray, 1997). Early growth and
tive interference of weeds with fruit trees yield of fruit trees in most reports have been
(Haynes, 1980). In studies of irrigation and lower in sod compared with herbicide-
nitrogen fertilization in orchards with differ- treated plots, and this has usually been
ent OFM systems, irrigation has often been attributed to water and nitrogen stress
more effective than nitrogen fertilization for (Hogue and Neilsen, 1987). In a recent OFM
promoting tree growth and yields (Hogue study where microsprinkler irrigation pro-
and Neilsen, 1987; Hipps et al., 1990). vided supplemental water during periods of
Experiments with stable isotope 15N-labelled inadequate rainfall, yields of newly planted
fertilizers have shown that soil nitrogen is apple trees after the fourth year were not sig-
more readily available to trees in herbicide- nificantly lower in sod than in herbicide
treated strips compared with turf grass, pre- treatments, and the best cumulative yields
sumably because grasses have a relatively occurred in post-emergence herbicide and
greater affinity for fertilizer nitrogen woodchip mulch plots, despite the presence
(Atkinson et al., 1979). However, drip-irri- of considerable weed populations during
gated apple trees in weed-free strips as nar- most of the year in those treatments (Plate
row as 0.3 m surrounded by turf grass have 13.2 and Fig. 13.3). These observations sug-
grown and yielded as well as non-irrigated gest that irrigation can reduce or compensate
trees with or without added nitrogen in for weed interference in established apple
wider weed-free strips (Broeshart and plantings.
225
200 Mulch
Fruit yield (kg per tree)

175 Post-herb
Pre-herb
150 Sod
125
100
75
50
25
0
1994 1995 1996 1997 1998 1999 2000 Cumulative
Fig. 13.3. Average and cumulative fruit yields over 7 years for trees in four OFM systems: hardwood-chip
mulch (Mulch); post-emergence (glyphosate) herbicide (Post-herb); residual pre-emergence herbicides (Pre-
herb); and a mowed sod grass (Sod). Repeated-measures analysis of variance (ANOVA) P values were 0.09
for treatments and 0.001 for time effects; data from an ongoing unpublished study of I.A. Merwin and T.
Steenhuis.
In flood- or furrow-irrigated orchards, knowledge and consideration of tree and

the movement and infiltration of water weed root physiology and site-specific
across and into the soil surface and the factors, as well as cost/benefit relationships
evapotranspiration of soil water reserves and consideration of long-term ecological
are affected substantially by OFM practices. processes that determine the sustainability of
Water evapotranspiration increased as orchard agroecosystems.
much as 40% in California orchards with
alleyway cover crops in comparison with
completely weed-free herbicide systems 13.2 Contemporary and Traditional
(Prichard et al., 1989). While surface flow is Roles for Orchard Ground Covers
more rapid in weed-free systems, the sedi-
ment loads in irrigation tail water are also Apples are often grown in cool humid
greater and water infiltration rates are regions of temperate latitudes, where pre-
reduced (Logan et al., 1987; NRC, 1993). cipitation occurs during much of the year. In
Soil-water consumption is greater in such regions, it is often necessary to apply
orchards with ground-cover vegetation and, pesticides or fertilizers, prune trees or har-
without irrigation, soil-water availability is vest fruit during inclement weather, when
reduced substantially, even in regions with machinery traffic over wet soils can lead to
frequent rainfall during the growing season compaction, rutting and erosion. To mini-
(Prichard et al., 1989; Hipps et al., 1990; mize such problems, perennial turf grasses
Merwin et al., 1994). are usually maintained between tree rows
In summary, the interactions among fruit (Hogue and Neilsen, 1987). The densely
trees, weeds or ground covers and soil and matted root and shoot systems of turf
water resources are affected substantially by ground covers help to protect soils from
OFM systems. Inadequate control of weed shearing and compression forces, reduce
competition for limiting resources can have mud and debris on fruit and equipment and
disastrous consequences for fruit growers. facilitate orchard management operations.
Although the complete eradication of weed In most apple orchards, permanent turf
competition is feasible with residual herbi- grass or seasonal cover crops are maintained
cides, it may not be necessary or desirable in the alleys, despite the added costs of
for economic and environmental reasons. mowing or cultivation and potential for
Integration of weed- and soil-management resource competition with the tree crop
tactics into comprehensive systems requires (Table 13.1).
308 I.A. Merwin
Table 13.1. Important characteristics of grass and legume ground covers maintained on the orchard
floor in apple-growing regions (from Heath et al., 1985; Skroch and Shribbs, 1986; Hogue and Neilsen,
1987; Prichard et al., 1989; Merwin and Stiles, 1994).
Nutrient use C:N

Ground-cover type (N–P–K kg year1) ratio Water use Establishment Vigour Durability
Red fescue 60–80–40 40 Low Very good Low High

(Festuca rubra L.)
Hard fescue 50–80–40 40 Low Good Low High
(Festuca duriuscula L.)
Tall fescue 50–60–40 50 High Very good High High
(Festuca arundinacea Schreb.)
Kentucky bluegrass 70–80–40 35 Moderate Good Medium Medium
(Poa pratensis L.)
Perennial ryegrass 60–80–40 30 Moderate Very good Medium Low
(Lolium perenne L.)
Annual ryegrass 50–70–40 40 Moderate Very good High Low
(Lolium multiflorum Lam.)
White clover 10–80–60 16 High Very good Medium High
(Trifolium repens L.)
Red clover 10–90–60 18 High Good High Medium
(Trifolium pratense L.)
The scenic beauty of many fruit-growing more limiting than labour costs, it can be
regions has made them important eco- advantageous to intercrop orchards with
tourism centres in Europe and elsewhere vegetables, other fruit crops (e.g. strawber-
around the world. In such regions, fruit ries beneath fruit trees), forage crops for
growers often benefit directly from agro- hay or pasture and sheep or cattle grazing.
tourism, offering pick-your-own sales or Fruit-yield reductions from intercrop
bed-and-breakfast lodging for visitors. The resource competition in such situations can
aesthetic and ecological aspects of OFM be offset by the extra income from addi-
systems are especially important in such tional crops or usages.
situations, and some growers plant attrac- In the past, orchards often served dual
tive cover crops or defer weed control dur- purposes for fruit growing and livestock
ing harvest or peak tourist seasons to make grazing, hence pasture was the original OFM
their orchards more pleasant to view, visit system in many regions (Morgan and
or harvest. Richards, 1993). With the advent of stringent
The high cost and limited availability of cosmetic and phytosanitary standards,
land suitable for fruit growing generate eco- which require routine pesticide applications
nomic pressures to use that land intensively, for fresh market apples, the use of orchards
hedging against price fluctuations or crop for livestock grazing has become less feasi-
failures by multiple land uses or cropping ble. Still, in parts of France, Spain and
systems. Polyculture – the simultaneous England where apples are grown primarily
cultivation of several interspersed crop for cider fermentation and the need for pesti-
species on a single field – is rare in commer- cide sprays is negligible, trees are high-bud-
cial orchards of Europe and North America, ded on robust rootstocks and grass/legume
because it increases labour requirements. mixtures are maintained throughout the
However, for diversified and subsistence orchard for livestock grazing beneath trees
farmers in regions where land costs are during much of the year.
Cover crops or crop rotations during depends on the soil fertility, availability of
orchard renovation and replanting can help irrigation and age of the trees (Broeshart and
to suppress problematic weed species and Keppel, 1985; Hipps et al., 1990; Merwin
soil-borne pathogens that cause root disease and Ray, 1997). In most situations, a single
and tree stunting (Mai and Abawi, 1981). combined application of pre- and post-emer-
Merwin and Stiles (1989) reported that pre- gence herbicides provides sufficient weed
plant cover crops of ‘Sparky’ marigold suppression in the tree row throughout the
(Tagetes patula L.) or ‘Saia’ oats (Avena year, making this the most economical OFM
sativa L.) suppressed lesion nematodes system (Merwin et al., 1995). Alternatively,
(Pratylenchus penetrans Cobb) and substan- several applications of post-emergence her-
tially increased the subsequent growth of bicides (e.g. glyphosate) have also been
apple seedlings in a New York orchard with shown to provide adequate control of weed
severe replant disease. In Washington competition in tree rows during the growing
orchard soils, Mazzola and Gu (2000) season, allowing a sparse ground cover to
observed substantial suppression of root dis- establish during the dormant season (see
ease and apple stunting, attributed to Plate 13.2) and providing some protection of
Rhizoctonia solani Sacc. AG5, following previ- the soil during the months when erosion and
ous crops of wheat (Triticum aestivum L.). nutrient leaching are greatest in humid tem-
Other cover-crop species, such as Sudan perate zones (Haynes, 1980; NRC, 1993).
grass (Sorghum sudanense Hitchc.) and mus- Observations of nitrate-nitrogen in drainage
tards (Brassica nigra Koch and Brassica juncea water from OFM plots in an established
Coss), have suppressed root pathogens of orchard with grass lanes between different
apple under laboratory conditions (Mayton tree-row treatments have indicated relatively
et al., 1996). Under field conditions in many low nitrate leaching in orchards compared
New York orchards, the response of newly with other agronomic crop systems
planted trees following purportedly disease- (Fig. 13.4), but nitrogen-leaching losses were
suppressive cover crops has been inconsis- usually greater in the herbicide treatments
tent and variable from site to site (Mai et al., relative to mulch or sod-grass plots (Merwin
1994; Merwin et al., 2000). Using ground et al., 1996).
covers or pre-plant cover crops as alterna- Soil-incorporated seasonal cover crops
tives to chemical soil treatments to suppress with herbicide strip systems are increasingly
soilborne pathogens of apple trees remains a being used in fruit-growing regions with
promising but unproved alternative (Pruyne mild, wet winters and long, dry, growing
et al., 1994; Biggs et al., 1997). seasons, such as California or the
Mediterranean area (Elmore et al., 1989;
Prichard et al., 1989). In this system, a weed-
13.3 Practical Options for OFM free strip is maintained in the tree row with
herbicides or cultivation, and the alleyways
A combination of herbicide applications in are seeded with annual grasses or a naturally
the tree row and mowed turf in the alley- occurring cover crop is allowed to establish
ways has become the norm among apple during the dormant season. This system
growers worldwide. In this system, a contin- facilitates irrigation and water infiltration
uous strip beneath the trees is treated with during the summer months, maintains soil
pre- or post-emergence herbicides to main- organic-matter residues, minimizes erosion
tain a more or less weed-free area. This mini- during the dormant season and ties up (or
mizes resource competition from weeds or fixes, in the case of legume cover crops)
ground covers by controlling their proximity nitrogen and other nutrients during the win-
to trees, and concentrates tree roots within ter when there is little uptake by the crop
the bare-soil strip, where they can proliferate (Perret et al., 1991; NRC, 1993). It has become
and utilize the soil volume and fertilizers or increasingly popular in regions where nitro-
irrigation more effectively (Atkinson, 1980). gen or phosphorus contamination of surface
The optimal weed-free area in this system and groundwaters are problematic (Wein-
310 I.A. Merwin
Drainage water nitrate-N (p.p.m.)

3.5
Mulch
3.0 Post-herb
Pre-herb
2.5
Sod
2.0
1.5
1.0
0.5
0.0
Spring Summer Autumn Winter Spring
1999 1999 1999 1999–2000 2000
Fig. 13.4. Seasonal average nitrate-nitrogen concentrations (p.p.m. or mg N l1) in drainage water samples
under four OFM systems in a mature apple orchard without nitrogen-fertilizer applications: hardwood-chip
mulch (Mulch); post-emergence glyphosate herbicide (Post-herb); residual pre-emergence herbicides (Pre-
herb); and a mowed sod grass (Sod). Data points are means of 12–36 water samples per season in each
treatment, from an ongoing study described in Merwin et al. (1996).
baum et al., 1992; Hornig and Bunemann, On poorly drained soils in humid regions,
1996a,b; Tagliavani et al., 1996a,b; Gut et al., prolonged water saturation of soil beneath
1997). In vineyards, this system has also been mulches can increase the incidence of root
shown to increase the quality of grapes and and crown rots caused by Phytophthora spp.,
the diversity and abundance of beneficial causing substantial tree mortality (Merwin et
arthropods (Remund et al., 1992), and similar al., 1992). Another disadvantage of mulches
benefits are likely in orchards. is their greater costs for establishment and
The mulch-strip turf-grass alleyway maintenance compared with sod or herbi-
system provides a practical non-chemical cide OFM systems (Merwin et al., 1995).
alternative to herbicide-strip systems. It Despite these problems, biomass mulches
involves applications of fabric, film or bio- can provide an effective non-chemical sys-
mass mulches along the tree row to suppress tem for reducing weed interference, conserv-
weeds, and a mowed turf-grass or seasonally ing soil moisture and enhancing soil fertility
cover-cropped alleyway. For certified ‘organic’ in orchards.
growers, mulches are a preferred OFM sys- Herbicide-treated bare-soil systems, in
tem. Biomass mulches provide substantial which surface vegetation is eliminated
quantities of organic matter, potassium, throughout the orchard, have provided the
nitrogen and trace elements to the soil and greatest long-term fruit yields in several
indirectly to trees (Haynes, 1980; Hogue and studies (Robinson and O’Kennedy, 1978;
Neilsen, 1987; Merwin and Stiles, 1994; Atkinson, 1980; Hipps et al., 1990) and may
Merwin et al., 1994). They increase soil-water be advantageous for irrigation or harvesting
retention and availability, keeping the soil of tree fruits and nuts in some regions
warmer in winter and cooler in summer, (Elmore et al., 1997). Long-term use of certain
which may be advantageous in some residual or post-emergence herbicides on the
regions. Weed species that thrive in cool, entire orchard floor has promoted the estab-
moist soils can eventually become a problem lishment of herbicide-resistant moss (Bryum
in biomass mulches, and may require argenteum L.) or liverwort (Marchantia poly-
supplemental control measures such as morpha L.) layers, which withstood traffic
spot-spraying with herbicides or manual and were apparently non-competitive with
weeding. Populations of voles (Microtus trees (Robinson and O’Kennedy, 1978;
spp.) and other rodents have increased Atkinson, 1980; Bastian, 1987; Merwin et al.,
under hay–straw, plastic film and fabric 1994). Cumulative yield increases of 20%
mulches, causing subsequent tree injury and have been reported where herbicides have
yield losses (Byers, 1984; Merwin et al., 1999). been used over the entire orchard floor, com-
pared with herbicide strip/turf alleyway herbaceous perennials. In many regions,

systems, but fruit quality (firmness, soluble woody perennials, such as poison ivy
solids and blush coloration) has often been (Toxicodendron radicans L.) and Virginia
reduced in comparison with herbicide strip creeper (Parthenocissus quinquefolia Planch.),
or overall turf-grass systems (Plate 13.1; also occur and pose serious problems if
Hogue and Neilsen, 1987). Without surface- allowed to proliferate beneath fruit trees.
vegetation residues to protect the soil from Such weeds must be rigorously suppressed
rutting, crusting and erosion, overall herbi- with herbicides or manual weeding at first
cide treatment or cultivation systems are appearance, and are usually held in check by
likely to cause long-term degradation of soil selective herbicides and routine mowing in
and water quality, and the trend in most the alleyways.
regions is to avoid year-round overall herbi- Each weed-control method selects over
cide systems in orchards (NRC, 1993). time certain ground-cover species with traits
Mechanical cultivation can be combined conferring resistance, tolerance or avoidance
with herbicide strips or sod alleyways to of that suppression tactic (Aldrich, 1984).
provide weed control during the growing With regular mowing over many years, a
season or to incorporate dormant-season complex mixture of herbaceous ground-cover
cover crops. In traditional plantings of large species develops in orchards, closely resem-
trees, it was often possible to plough or bling cattle pastures in cool humid regions. In
harrow in several directions parallel and a New York study on a silty loam soil, inten-
perpendicular to tree rows, providing weed sive sampling of a regularly mowed orchard
control over most of the orchard floor. The floor 1 year after renovation and seeding a
practice of clean cultivation was widely rec- turf-grass mixture of perennial ryegrass
ommended and adopted during the early (Lolium perenne L.) and red fescue (Festuca
1900s, but has been discontinued in most rubra L.) revealed that more than 72 species
regions now for several reasons. Soil cultiva- were present within a 0.9 ha site, with ten
tion inevitably damages some tree roots, but, predominant species each comprising at least
in deep soils or in regions where the upper 1% of total ground cover in all 50 sampled
soil layer is too hot during the summer for plots (Merwin, 1990). After 4 years under dif-
extensive proliferation of apple roots, it does ferent OFM systems in that study (mowing,
increase tree growth and productivity com- mulching, monthly cultivation and selective
pared with traditional sod orchards (Haynes, pre-emergence or broad-spectrum post-emer-
1980). Continuous cultivation also reduces gence herbicide treatments), distinct species
soil organic matter and structural integrity mixtures had developed in each system.
(Fig. 13.1), increases the potential for soil ero- Herbicide-tolerant ground-ivy (Glechoma heder-
sion due to exposure to water and wind and acea L.), pigweed (Amaranthus retroflexus L.)
eventually leads to compaction and reduced and milkweed (Asclepias syriaca L.) became
soil fertility compared with no-till herbicide dominant in pre-emergence herbicide plots.
or sod systems (NRC, 1993; Merwin et al., Mosses and weeds, such as dandelions
1994). For these reasons, cultivation is often (Taraxacum officinale Weber) and groundsel
used in rotation with dormant-season cover (Senecio vulgaris L.), with seeds dispersed
crops, where the cover crops help to re- readily during midsummer, infested sparsely
plenish soil organic-matter residues, reduce the post-emergence herbicide (glyphosate)
erosion and retain nutrients. plots. In straw-mulch plots, weeds such as
milkweed, couch grass (Elytrigia repens L.)
and curly dock (Rumex crispus L.), which
13.4 Ground-cover Vegetation Species, thrive in moist, nutrient-rich soils and pro-
Types and Characteristics duce persistent deep rhizomes or tap roots,
became problematic after 4 years. In the fre-
The ground-cover vegetation in most quently cultivated (rotavated) plots, couch
orchards is a complex mixture of three grass, wild carrot (Daucus carota L.) and crab-
types: grasses, legumes and broad-leaved grass (Digitaria sanguinalis (L.) Scop.) became
312 I.A. Merwin
the dominant weeds. Elmore et al. (1989) and spp.) or alfalfa (Medicago sativa L.), are often
Skroch et al. (1975) have reported similar grown in orchards for their soil-nitrogen con-
weed-species selection trends over time; each tributions. Turf grasses often benefit from
method of OFM selects for certain types of interspersed clovers, with the combination
ground-cover species with distinct ecologi- providing a more durable ground cover
cal and management characteristics. Rotation (Heath et al., 1985). In California, strawberry
among different weed-management systems and subclovers were reportedly useful
is an appropriate tactic for minimizing the orchard ground covers that provided soil
adverse effects of system-related weed- protection and nitrogen inputs during the
selection processes. winter months and became dormant during
Various types of grasses are seeded or the summer growing season (Elmore et al.,
occur naturally in orchards, including 1989). Until recently, clovers were recom-
species with C-3 and C-4 photosynthetic mended as ground covers in apple orchards
traits, cool- or hot-season growth phases and of the eastern USA, but for numerous rea-
erect, clumped or sprawling growth habits. sons legumes may not be desirable in apple
A detailed description of common grasses is orchards (Hogue and Neilsen, 1987). The
beyond the scope of this chapter, but many uncontrolled release of nitrogen from legume
grasses are excellent ground covers for roots and residues in late summer could
orchards (Table 13.1). Turf grasses, especially reduce tree winter-hardiness and fruit
those of prostrate growth habit that spread quality, but there is little evidence that this
by runners and/or rhizomes, are quite resis- actually occurs. Fruit damage from Lygus
tant to machinery and pedestrian traffic. plant bugs (Lygus lineolaris L.) and the
Most orchard grasses have a high affinity for vectoring of graft-union necrosis virus into
soil nitrogen and water (Hogue and Neilsen, apple rootstocks are exacerbated with
1987; Merwin et al., 1994), with dense but rel- legume ground covers. The risk of honey bee
atively shallow root systems compared with poisoning increases greatly when insecti-
fruit trees. Grasses do not usually serve as cides are applied in orchards where legumes
alternative hosts for virus diseases and are flowering, and the relatively deep-rooted
arthropod pests affecting fruit crops, and growth of most legumes increases their
they respond positively to routine mowing, competition with trees for soil water and
forming a dense layer of thatch and turf that nutrients (Merwin and Stiles, 1994). For
resists invasion by other herbaceous weeds. these reasons, apple growers often suppress
Low-stature cool-season grasses, such as red legumes, using broad-leaved-selective herbi-
fescue or hard fescue (Festuca duriuscula L.) cides such as 2,4-D (2,4-dichlorophenoxy-
are preferred alley ground covers in many acetic acid).
apple-growing regions, because they grow A multitude of broad-leaved herbaceous
less vigorously and require less mowing dur- perennials occur in orchards, and most are
ing the midsummer season when nutrient treated as weeds and suppressed with her-
and water stress are maximal for fruit trees. bicides or cultivation. Recent studies indi-
Seeding annual grasses, such as Italian rye- cate that flowering ground covers that
grass (Lolium multiflorum Lam.), may be use- provide nectar, pollen or habitat for benefi-
ful in late summer to provide moderate cial arthropods can increase the biological
nutrient stress, which improves fruit quality, control of pests in fruit trees and vines
reduces pruning costs and nutrient leaching (Remund et al., 1992; Bugg and
and protects soil during the winter months Waddington, 1994; Gut et al., 1997). There is
(Tagliavani et al., 1996a; Mantinger and ample evidence that beneficial arthropod
Gasser, 1997). abundance and diversity are greater in
Legumes, such as white and red clovers orchards with flowering annual and peren-
(Trifolium repens L. and Trifolium pratense nial ground covers (Altieri and Schmidt,
L.), subterranean and strawberry clovers 1986), but little evidence that increasing
(Trifolium subterraneum L. and Trifolium beneficial populations in the understorey
fragiferum L., respectively), and vetches (Vicia results in significant levels of pest control
within the fruit and foliage of apple (Brown 13.5 Weed-control Methods and
and Glenn, 1999). Further work is needed to Equipment
quantify the trade-offs between potential
biocontrol advantages and probable disad- 13.5.1 Herbicide applications
vantages, such as management complexity,
resource competition between flowering Weed resistance to certain herbicides is a
ground covers and fruit trees and the risks known problem in agronomic crops and a
for honey bees and other beneficial insects serious concern in fruit production, but at pre-
foraging on flowers during insecticide sent there are effective herbicides available to
applications. Research in North Carolina control the major weed species in apple
and California has indicated that some orchards (Derr, 2001). Tank mixing of selective
ground-cover species are relatively non- herbicides and precisely directed spray appli-
competitive with fruit trees and can be pro- cations enable growers to obtain sufficient
moted by selective management practices control of weed competition in most circum-
(Skroch and Shribbs, 1986; Elmore et al., stances (Tables 13.2 and 13.3). Various types
1989). Fruit growers around the world are and designs of herbicide sprayers are avail-
evaluating reduced-tillage and seasonal able, including shielded-boom sprayers, ultra-
cover-crop systems as alternatives to clean- low-volume sprayers, wipers and backpack
cultivation and residual-herbicide OFM. spot sprayers. Equipment is also available for
Table 13.2. Residual pre-emergence herbicides used in apple orchards. This list is for general reference
only, and does not imply discrimination or endorsement by the author.
Dichlobenil For hard-to-kill perennial weeds such as couch grass, nutsedge (Cyperus esculentus
L.) and reduction of field bindweed (Convolvulus arvensis L.). Also effective for the
control of annuals. Should be lightly incorporated in arid climates or applied in the
rainy season for best results
Diuron For annual broad-leaved plants and grasses or in combination with bromacil or other
pre-emergence selective materials in some deciduous fruit crops
Napropamide For annual grass control and in combination with other selective herbicides for broad-
leaved plant control on young trees, because of its positional selectivity (surface
adsorption and retention) above the root zone. If mixed down into soil, severe tree
injury can occur
Norflurazon For long residual annual grass and some broad-leaved-plant control. Sometimes used
in combination with other selective herbicides
Oryzalin For annual grass and some broad-leaved-plant control. Often used with other products
to broaden spectrum of combinations. Applied around young trees and vines because
of its relative safety for these plants due to positional selectivity
Oxyfluorfen For a broad-spectrum broad-leaved-plant control material, often in combination with
one of the grass-control herbicides, such as oryzalin, pendimethalin or napropamide.
When used on young trees or vines, care should be taken to keep the spray off the
foliage, or contact injury can result
Pendimethalin For annual grass and some broad-leaved-plant control on young trees, often combined
with other herbicides to enhance the long-term residual grass control
Pronamide For annual and some perennial grass control, particularly couch grass
Simazine For broad-spectrum control of annual weeds, especially effective on annual broad-
leaved weeds and combined with other grass-control materials. Its long residual
activity has made it a popular herbicide. Has been found in groundwater after long-term
use in some locations
Terbacil For long-lasting residual control of annual grasses and broad-leaved weeds in
orchards
Trifluralin For pre-plant control of grasses and some broad-leaved weeds. Applied as a continu-
ous band under the soil surface, effective for the suppression of field bindweed in
young plantings
314 I.A. Merwin
Table 13.3. Post-emergence non-residual herbicides often used in apple orchards. This list is for general
reference only, and does not imply discrimination or endorsement by the author.
Fluazifop-P-butyl Used for selective control of most annual and perennial grasses
Glyphosate For selective control of most grass and broad-leaved species and woody weeds.
Injury will occur if sprayed on to leaves or green bark of trees
Glufosinate Primarily effective on annual grass and broad-leaved species, with less control of
perennials
Oxyfluorfen For control of young broad-leaved weeds, or in combination with other post-
emergence herbicides
Paraquat A non-selective contact herbicide used to control grass and broad-leaved annual
seedlings
Sethoxydim A selective herbicide for the control of most grasses, ineffective on broad-leaved
weeds
Sulfosate Non-selective herbicide for control of annual and perennial grasses and broad-leaved
weeds. Selectivity based upon non-permeable woody outer bark of mature trees and
vines
2,4-D Used for selective control of most broad-leaved weeds. Care must be taken to
minimize chance of drift on to leaves of crop plants, especially during warm weather
applying granular herbicides. In general, her- Backpack sprayers are useful where sparse or
bicides should be applied in a sufficient vol- scattered weeds require spot treatment, but
ume of water to wet a substantial portion of drift and inadvertent damage to trees is more
the surface area being treated, with minimal likely with backpack applications.
runoff. This often requires about 200–400 l of
spray material per treated hectare, though
substantially less may be needed with wiper 13.5.2 Between-row cultivators
or ultra-low-volume equipment. The optimal
volume of water also varies with the herbi- Various designs of rotavators and disc, brush,
cide being applied. For example, the effec- tine, rotary-hoe and chisel-tooth harrows are
tiveness of glyphosate is often greater with available for orchards. Mechanical cultivation
lower volumes of water, while that of usually provides only temporary weed sup-
paraquat is improved by increasing the vol- pression. Weed regrowth occurs after several
ume of water applied. weeks or months, depending upon ground-
Sprayer application pressure should be cover species mixtures and water availability.
kept low enough to minimize spray atomiza- As many as six cultivations per season may
tion in the nozzles and drift, but high enough be necessary to control weeds throughout the
to produce an even spray pattern in the noz- growing season, and perennial weed prob-
zles and across the desired boom width. All lems often increase when cultivation is the
sprayers should be carefully calibrated. sole weed-control method.
Overdoses of herbicides can seriously dam-
age fruit trees, while insufficient dosages
may not provide effective weed control. 13.5.3 Within-row cultivators
Shielded-boom herbicide sprayers, with
shrouds or curtains between the crop and the Many types of in-row cultivators are avail-
nozzles, provide an extra margin of safety by able and more are being developed and uti-
minimizing drift, especially under windy con- lized as alternatives to chemical herbicides.
ditions (Plate 13.3). Wipers are used to selec- Available equipment includes rotavators that
tively apply herbicide to one portion of the swing in and out of the row; horizontal cir-
ground-cover vegetation, with minimal chance cular tillers that scuff soil out of tree rows;
for drift or penetration to the crop or desirable flat blades that cut under weeds; and the tra-
species in the ground-cover vegetation. ditional hoe-plough, which turns a furrow
within the row and can be retracted in pass- ngs, due to problems such as air pollution,
ing by trees. All of these implements work fire hazard and damage to the trunks and
close to trees and can damage low-slung foliage of the crop. Preliminary work with
branches, trunks and shallow roots if in-row propane burners in apple orchards
improperly adjusted or operated. suggests that crop damage and fuel con-
sumption can be minimized if the heat
source is shrouded (Merwin and Ray, 1994;
13.5.4 Mowing equipment Plate 13.5). Flame or steam weed suppres-
sion is more short-term than contact herbi-
Mowing is primarily done in the row mid- cides, such as paraquat, but may be practical
dles of no-till orchards in older orchards in for organic and other growers who cannot
humid growing regions. There are mowers use chemical methods of weed control.
that cut within the row on trees or vines
planted on level grades, but most work
poorly in raised-bed or berm plantings. 13.6 Epilogue
Available implements include rotary, flail
and sickle-bar mowers. Some flail mowers There are many different systems for manag-
are designed also to grind up tree prunings ing surface vegetation and soil resources in
left on the ground in row middles, provid- orchards, and each has its strengths and
ing a thin mulch layer and retaining nutri- weaknesses. No single OFM system is best
ents in the orchard. Flail and rotary mowers for all situations. Their relative merits are
with side-delivery chutes can be operated to determined by soil and climate type, orchard
place the clippings as a mulch within tree age and design, grower preferences and
rows, helping to move nutrients from the resources, labour and capital availability,
sod to the tree row and suppressing weed market incentives or ‘green’ labels, such as
growth. Adjustable-width low-profile mow- organic or integrated fruit production, and
ers are especially useful for season-long local regulations concerning soil and water
management of vegetative ground covers quality. Moderate stress from managed weed
within the tree row, and they facilitate close competition at certain times of tree and fruit
mowing beneath trees with minimal dam- development may be preferable to total
age to heavily cropped branches at harvest weed eradication that leaves the orchard soil
time (Plate 13.4). without protective ground cover and prone
to erosion and nutrient loss during much of
the year. Ecological, environmental and eco-
13.5.5 Flame or steam weed suppression nomical factors must all be considered in
developing OFM systems that improve
Relatively little work has been done with orchard efficiency, enhance fruit quality and
flame or steam weed control in fruit planti- help to sustain vital natural resources.
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14 Pruning and Training Physiology
David C. Ferree1 and James R. Schupp2

1Department of Horticulture and Crop Science, Ohio Agricultural Research and
Development Center, Ohio State University, Wooster, Ohio, USA;
2Department of Horticultural Sciences, Cornell University, Highland, New York, USA

14.2 Pruning 320
14.2.1 Effects on growth 320
14.2.2 Effects on flowering 322
14.2.3 Effects on fruit 322
14.2.4 Effects on mineral nutrients 322
14.3 Types of Pruning Cuts 322
14.3.1 Thinning cuts 323
14.3.2 Heading cuts 324
14.4 Time of Pruning 324
14.5 Pruning Trees of Different Ages 325
14.5.1 Young trees 325
14.5.2 Fruiting trees 327
14.5.3 Old or declining trees 330
14.6 Summer Pruning 331
14.6.1 Effects on physiological processes 331
14.6.2 Effects on fruits 331
14.6.4 Practical implications 333
14.7 Root Pruning 334
14.7.1 Effects on physiological processes 334
14.7.3 Effects on fruiting 336
14.7.4 Practical implications 336
14.8 Training 337
14.8.1 Bending 337
14.8.2 Phloem interruption 339
14.8.3 Notching 341
14.9 Conclusion 341

320 D.C. Ferree and J.R. Schupp
14.1 Introduction of this chapter is to provide the physiological

basis for these cultural techniques so that
Pruning and training are almost always they can be used to best advantage.
mentioned together, but there is a difference
between them and each has a clear function.
Pruning has been defined as the art and 14.2 Pruning
science of cutting away a portion of the plant
for horticultural purposes. Pruning can be 14.2.1 Effects on growth
used to improve tree shape, to influence its
growth, flowering and fruitfulness, to improve Although the practice of pruning goes back
fruit quality, to repair injury, to contain the several thousand years, the science of prun-
plant and to encourage light and spray ing originated in the early 20th century, with
penetration. Training, in contrast, refers to experimental data revealing several general
the direction of tree growth or form and the influences. The first of these is that pruning
development of the structural framework of is a dwarfing process and the more severely
the tree. Pruning is only one of the tech- a plant is pruned, the greater the degree of
niques used in training. Although some dwarfing. Removal of the branch not only
training may be necessary after the tree removes stored carbohydrate and nitrogen
comes into production, training is mostly reserves, but also reduces potential leaf sur-
confined to the period when the tree is face and growing points as well. Obvious
becoming established. The ultimate goal of reductions in number of growing points, tree
both pruning and training is to improve light height and spread and leaf area can easily be
distribution so that as much of the tree observed on pruned versus non-pruned
canopy as possible maintains production of plants. Although not as obvious, pruning
high-quality fruit. Forshey et al. (1992) pre- correspondingly reduces root growth, and
sent a comprehensive treatise on pruning new root growth will be delayed until shoot
and training, including an extensive refer- growth in response to pruning occurs.
ence list. A comprehensive literature review There is a fundamental equilibrium
will not be included in this chapter but refer- between the above- and below-ground
ences will be cited to illustrate key points. components of an apple tree. Removal of a
Apple trees that are not trained and portion of either the top or root system slows
pruned become very large (Plate 14.1) and growth of the other until the balance is re-
develop fine branching at the tree periphery, established (Taylor and Ferree, 1981; Geisler
which becomes very dense and shades the and Ferree, 1984a). This equilibrium is proba-
tree interior. Fruit size, colour and sugar con- bly achieved not only through loss of reserves
centration decline on shaded spurs and, as in the tissue removed, but also through
light becomes still more limiting, the spurs changes in the functional equilibrium
themselves become weak and unproductive, between the root and the top and the balance
eventually dying. As the trees age, a larger of hormones produced in meristematic
and larger portion of the tree canopy volume regions in the leaves, shoot tips and root tips.
becomes unproductive. As the unproductive Shoot growth is dependent upon water
area of the canopy expands, it becomes a and mineral nutrient uptake by the root sys-
larger sink for photosynthates and thus tem, while roots depend upon the above-
decreases tree efficiency. The dense shading ground portion of the tree for carbohydrates.
encourages insects and diseases due to high Pruning away a portion of either of these
humidity, slow drying conditions and poor plant parts reduces its capacity to perform its
spray penetration. Thus, although pruning function; thus growth in other parts of the
and training are expensive cultural practices, plant is reduced and energy is redirected to
economic fruit production is not possible regenerate the missing component. This con-
without the improvement in light distribu- cept, known as functional equilibrium,
tion these techniques provide to maintain seems to be moderated by plant hormones
tree efficiency and fruit quality. The purpose (Richards and Rowe, 1977).
Pruning and Training Physiology 321
Grochowska et al. (1984) have shown that work of Verner (1955) illustrated that when
pruning the top during the dormant season four or more shoots were above a new
increases the supply of cytokinins from the branch, the angle of that new branch was
roots, measured as increased concentration in 50–90° (i.e. close to horizontal). When there
the remaining above-ground tissue. These were no shoots above the new branch, the
increased hormone levels are probably respon- branch angle was in the very narrow range
sible for stimulating cell division and ulti- of 0–40° (i.e. very upright). Removal of the
mately shoot growth, which, in turn, promotes terminal bud results in an increase in the
auxin production in the shoot tips and gib- growth of the subtending laterals, and
berellin production in the new, unfolding shoots that would have been spurs become
leaves. The increases in concentration of these vegetative shoots. Removal of apical domi-
three classes of hormones found in shoot tissue nance not only changes which shoots grow
in the early part of the growing season follow- and their vigour, but the resulting shoots are
ing dormant pruning not only increase growth more upright with narrower crotch angles.
but also alter the pattern of growth. Removal Thus, tree form is changed into a more
of terminal buds and their auxin results in upright, narrow shape.
enhanced growth of laterals of latent and Even though pruning dwarfs the whole
existing flower buds. The more upright the plant or limb, an invigorating growth
branch that is pruned, the more vigorous will response occurs in the immediate area of the
be the growth that responds to the removal of cut. This invigorating effect decreases as the
apical dominance (Myers and Ferree, 1983b). distance from the cut increases. This is an
Normally a terminal bud makes the most important general concept to understand,
growth in an unpruned, upright branch, and because, if the purpose of pruning is to
each subsequent subtending lateral makes invigorate and stimulate growth, as is often
progressively less growth, until some form the case in areas of the canopy that are spur-
spurs, with the most basal buds not grow- bound, more response is generated by many
ing. This growth form is probably the result small cuts than by a few large ones (Ferree
of cumulative auxin concentrations pro- and Forshey, 1988). Generally, many small
duced from each of the buds above causing cuts stimulate more shoot growth, reduce
a reduction in growth in the shoot originat- fruiting to a greater degree and alter tree
ing at lower levels (Fig. 14.1). The classic form more than when a few large cuts are
made. Larger cuts result in fewer, more vig-
orous and more localized regrowth. Another
manifestation of the localized influence can
be observed in the injury that occurs when
trees are pruned close to a sudden drop in
winter temperature. The freezing injury
results in death of the bark and phloem
tissue and is most obvious close to pruning
cuts, at the tree base and in limb crotches
on pruned trees compared with unpruned
adjacent trees (Rollins et al., 1962; D.C.
Pruning cut Ferree, personal observation). This localized
response occurs even though the tree appears
to be completely dormant.
One of the primary functions of pruning
is to develop a framework of branches that
are strong, likely to consistently carry a
good crop of premium-quality fruit and be
Fig. 14.1. Natural growth pattern of apple (left) and less prone to environmental extremes. To
change in the pattern caused by a heading cut achieve this goal the crotch angle, which
(right). forms very early in the first year of the
development of a branch, must be wide and pruned trees because the supply of reserves
strong. Narrow crotches cause bark inclu- available to remaining blossoms is
sions, are weak, split easily and are more sus- increased. Cell division early in the season is
ceptible to cold injury. Wide crotch angles are greater in areas of the canopy that have high
formed naturally in very vigorous-growing light and large spur-leaf areas. Later in the
trees that have very active meristematic season during cell expansion, shoot leaves
areas that produce large quantities of auxin supply the fruit. Fruit size is closely related
(Verner, 1955). to spur-leaf area and well-illuminated
shoots (Ferree and Palmer, 1982; Rom and
Ferree, 1986). Pruning is the cultural practice
14.2.2 Effects on flowering most influential in establishing a canopy
light environment that promotes high spur
Another general principle is that pruning quality and an even distribution of shoots
delays the flowering and fruit bearing of through the canopy.
young trees and decreases fruiting in
mature trees. Forshey et al. (1992) state that
this is the result of the following three fac- 14.2.4 Effects on mineral nutrients
tors: (i) the removal of effective bearing sur-
face; (ii) the stimulation of vegetative Another indirect effect of pruning is the
growth from growing points that were for- influence of pruning on the mineral content
mer or potential fruiting spurs; and (iii) of the tree and fruit. The decrease in number
direct competition between vigorous shoot of fruits caused by pruning is associated
growth and fruit set. They indicate that the with an increase in leaf nitrogen, potassium
reduction in cropping is proportionately and phosphorus and a reduction in fruit
much greater than the reduction in total concentrations of magnesium and calcium.
growth. Pruning young trees tends to force These changes are associated with the com-
growth of long succulent shoots, which con- petition of the fruits and vigorous shoots for
tinue to grow late in the season. Thus, car- the elements and with the increase in fruit
bohydrates do not accumulate for size caused by dormant pruning. Since cal-
flower-bud initiation, but are used in pro- cium is so important to fruit storage poten-
ducing the vigorous vegetative growth tial and disorders such as bitter pit, the
(Mika, 1975). Elfving (1990) demonstrated reduction in fruit calcium caused by pruning
that heading back pruning in a single year is a particular concern. Since vigorous shoots
significantly reduced tree size and yield and are a stronger sink for calcium than the fruit,
the effect remained evident 5 years after the dormant pruning reduces fruit calcium and
pruning. The more severe the pruning, the the more severe the pruning, the greater the
greater is the decrease in fruiting (Elfving reduction. Dormant pruning also increases
and Forshey, 1976). The negative influence fruit size, which leads to a dilution effect on
of pruning on fruiting is particularly impor- calcium concentration. In contrast, summer
tant in intensively planted modern orchards pruning, which removes vegetative shoots,
that are dependent on early cropping to can increase fruit calcium (Preston and
help control growth. Perrin, 1974).
14.2.3 Effects on fruit 14.3 Types of Pruning Cuts
One of the primary reasons for annual prun- All pruning cuts can be classed into two
ing is to increase fruit size. The effect on types (thinning out and heading back) and
fruit size is not a direct effect of pruning, but each type of cut produces a different physio-
results from the reduced number of fruit logical response. A thinning cut (Fig. 14.2)
and improved light conditions in the removes an entire shoot, spur, branch or
canopy. Generally fruit set is higher on limb, while a heading cut (Fig. 14.3) removes
Dormant Unpruned Pruned – thinning cuts

Fig. 14.2. Change in growth of an apple tree that received two dormant thinning cuts in year 1 (left-hand
figure). Growth response at the end of year 2 is shown in the right-hand figure. The response for an
unpruned tree is shown in the centre figure. In the latter case, main pruning cuts are needed in the second
dormant season to restore a desired tree form but note that the advantages of lateral branch development
and uniform fruit distribution have been lost.
Dormant Unpruned Pruned – heading cuts

Fig. 14.3. Change in growth of an apple tree that received three heading-back pruning cuts in year 1
(left-hand figure). Growth response at the end of year 2 is shown in the right-hand figure where lateral
branch development is strong and vigorous upright growth is apparent. The unpruned tree (centre figure) has
a more spreading growth form with less lateral branching.
only a portion of a shoot or limb, leaving remove unfruitful wood that is upright and
another portion from which new growth can vegetative or pendant and weak.
develop. A comparison of the various Generally, thinning cuts improve light dis-
responses from the two types of cuts is sum- tribution in the tree, thus increasing flower
marized in Table 14.1. initiation, fruit set and spur growth in lower
parts of the canopy. The effect of thinning cuts
on fruiting is limited to the amount of poten-
14.3.1 Thinning cuts tial bearing surface removed. The ratio of ter-
minal to lateral buds is largely undisturbed
Thinning cuts are primarily used to alleviate and, by removing branches at their point of
crowding and shading and generally origin, only weaker latent buds are left to
improve light penetration into the canopy grow; thus, thinning out does not increase
(Fig. 14.2). Thinning cuts also function to shoot growth as much as heading back.
Table 14.1. Relative influence of thinning and heading pruning cuts on apple trees.
Response Thinning cuts Heading cuts
Apical dominance Little effect on remaining growth Terminal removed, buds released
Shoot formation Fewer vegetative shoots Increased number and length
Spur formation Increased Decreased
Tree shape Limited effect Dramatic effect, e.g. hedging
Tree growth habit Natural tall rangy Compact and upright habit
Fruitfulness Limited effect Significant decrease
Carbohydrate reserves Limited effect Dramatic reduction
Nutrition elements Limited effect Decrease calcium, magnesium,
increase nitrogen
Local stimulus Encourage few shoots Encourage many shoots
The thinning-out cut should be just above growth and fruiting lasting 5 years.
the small ridge at the base of a branch, called Heading cuts stiffen limbs and care should
the collar. Cuts made at this location make be exercised in the use of these cuts on
the smallest wound, heal the fastest and are upright branches of young trees because
less susceptible to winter injury. Leaving a some type of physical limb bending will be
stub above the ridge or cutting too close and required to return these branches to a bal-
removing the ridge slows healing and pro- anced fruiting condition.
vides more potential for disease or insect Each type of cut has a role in creating and
damage (Shigo, 1990). An exception to this managing an efficient tree with a good bal-
rule is the Dutch cut, a bevelled stub that is ance between vegetative vigour and fruiting.
left for stimulating a renewal limb with cer- Newly planted trees are headed to stimulate
tain training systems (see Chapter 15). the production of strong lateral branches
needed to form scaffolds. Thinning cuts are
necessary on young trees to develop the
14.3.2 Heading cuts desired primary branch framework with
minimal effect on fruiting. As trees reach full
Heading cuts (Fig. 14.3) locally stimulate size, thinning cuts maintain the adequate
more vegetative growth, which tends to be light and spray penetration needed to keep
vigorous and shades the tree interior, result- the whole canopy productive, while fruit
ing in fewer fruiting sites and less flower size and quality are improved. Cultivars
initiation than thinning cuts. Since using this such as spur-type ‘Delicious’ often form
type of cut removes many terminal buds, insufficient numbers of laterals and may
apical dominance and the hormone balance benefit from heading cuts to induce laterals
is disturbed more than with thinning cuts. where they are needed. In older trees, when
Heading increases both shoot number and portions of the canopy become weak or spur-
length, but the effect on length is determi- bound, heading cuts can encourage shoot
nate. The younger the wood and the more growth and improve fruit size.
upright the growth headed, the greater the
effect. Heading 1-year-old wood results in a
strong localized response, releasing the 14.4 Time of Pruning
three to four buds below the cut to develop
into vigorous, upright shoots with narrow Most pruning on apple trees is done when
crotches. Heading causes many shoots that they are dormant. In areas that can experi-
would have been potential spurs or fruiting ence very low temperatures in winter, time
shoots to form vigorous vegetative growth. of pruning is an important consideration
Elfving (1990) has shown that heading cuts because it increases susceptibility to low-
on young trees have negative effects on temperature injury (winter injury). Sensi-
tivity to cold injury after pruning is greatest 14.5 Pruning Trees of Different Ages
in the tissue in close proximity to the prun-
ing cuts. The exact temperature required to 14.5.1 Young trees
induce winter injury varies with many fac-
tors, such as previous exposure to low tem- Most apple orchards are trained to some
peratures, previous crop, cultivar, rootstock modified form of a central leader system,
and tree age. Winter injury may occur at resulting in a cone-shaped canopy. In this
low temperatures in the range of 23 tree form, the central stem, or leader, is
to 49°C, depending upon the inherent maintained in an upright orientation, while
hardiness of the tree and these contributing limbs that form on the sides are trained at an
factors. Pruning within 2 weeks prior to the angle to form the primary scaffolds.
low-temperature event will increase the Additional secondary limbs form on both the
sensitivity of the tree to cold injury, with the leader and the scaffolds and are maintained
greatest loss of hardiness occurring within to provide a bearing surface. The purpose of
the first 48 h (Rollins et al., 1962). The loss in this chapter is not to provide a ‘how-to’ trea-
hardiness is then gradually regained. Thus, tise on pruning trees trained to specific sys-
the risk of causing winter injury by pruning tems, but to cover the principles and
can be reduced by suspending pruning physiological basis for practical application
work when severe cold weather is forecast for a generalized central-leader system. The
(see Chapter 10). specific alternative pruning practices for
To reduce the risk further in areas prone young trees in specific training systems will
to winter injury, pruning should be delayed, be covered in Chapter 15.
where practical, until after the coldest months Trees that come from the nursery have
(January and February in the northern lost a significant amount of their root sys-
hemisphere). If pruning must be started ear- tem and thus the top needs to be reduced
lier, it is best to prune the oldest trees and to create a balance so that sufficient growth
most cold-tolerant cultivars first and to delay occurs to form the permanent scaffolds. If
pruning the youngest trees and most cold- trees arrive as unbranched whips, they are
sensitive cultivars. usually headed back 60–90 cm above the
When pruning is delayed until growth ground, depending on which training sys-
starts, the tree responds differently. Delay tem is used. Cultivars such as spur-type
much beyond bloom devitalizes growth and ‘Delicious’ do not branch easily and will
may interfere with the development of only produce shoots within 5 cm of the
flower buds for the next year’s crop. Pruning heading cut, while cultivars such as
cuts at the time vegetative growth is begin- ‘Golden Delicious’ will produce more later-
ning result in an increase in the number of als, which are distributed 10–15 cm below
buds that break and buds further from the the cut (Ferree, 1978). Cultivars that have a
cut tend to break more when compared with terminal bearing habit should be headed
dormant pruning. This response is probably high, as these cultivars tend to produce
due to the removal of apical dominance since weaker shoots from the lower buds, plus
the meristematic regions in the growing the terminal bearing habit causes the
buds are primary sources of auxin. Generally branches to bend, which can cause them to
the breaks that occur from delayed pruning interfere with herbicide applications and
are less vigorous and may be very desirable mowing operations if they originate too
on cultivars that tend to have blind wood low on the trunk.
(previous-season wood with no lateral If feathered trees (see Chapter 6) are
growth), such as ‘Tydeman’s Red’ or ‘Rome planted, shoots to form the primary scaffolds
Beauty’. Delaying dormant pruning until are already in place and, generally, the only
bloom can also be used to lessen the cuts made on these trees are to remove any
regrowth in overly vigorous trees; however, spurs or shoots below 45 cm above the
delayed pruning should not be used on trees ground, as these interfere with weed control
with less than optimum vigour. and will produce few marketable fruit.
During the first growing season, leader of the remaining leader shoot. Cultivars such
and scaffold selection are very important as spur-type ‘Delicious’ may not produce
and should be done early so that the tree sufficient shoots to use this method of leader
directs its growth as much as possible into selection. In this instance, the shoots with
tissues that are to be retained. Each growing upright growth and very narrow crotch
shoot produces auxin, which moves down angles, which would be removed in the
the tree and affects the growth and crotch above procedure, are retained. A spring
angle of shoots below. The uppermost shoots clothes-peg is clamped around the leader
grow vertically because of an absence of just above the young shoot when it is 5 cm
auxin during and after bud break. The prolong, and it is forced to grow horizontal
gressively increasing auxin concentration below the clothes-peg (Fig. 14.4). This shoot
further down the trunk stimulates lower will ultimately curve upward, but only after
shoots to develop wider crotch angles and a wide, strong, crotch angle has been estab-
this effect is complete by the time the shoots lished. The growth of these shoots is reduced
are 5–10 cm long (Verner, 1955). This natural 15–20% by the bending process further
pattern can be used to advantage by select- establishing the dominance of the leader. An
ing the leader when the shoots are 10 cm alternative to the clothes-pegs is to use
long, removing the two or three upright pointed toothpicks or thin wire to force the
shoots that are growing just below the leader new shoot to grow laterally. These practices
and leaving the lower shoots with wider must be carried out before the base of the
crotch angles to develop as scaffolds. Since shoot lignifies. It takes about 1 min per tree
the tissues in question are succulent, care to install the clothes-pegs and about 15 s to
must be exercised to avoid injury to the vas- remove them 3–4 weeks later to avoid
cular tissue, as this can decrease the growth girdling the leader.
Headed, Headed, Headed,

untreated clothes-pegs in place subsequent form following
use of clothes-pegs
Fig. 14.4. Influence of clothes-peg spreading on growth of a newly planted unbranched apple tree that has
been headed.
The most important principle to remem- Vigour varies throughout the canopy, with
ber on young trees is to prune them as little greatest vigour in the top and periphery. This
as possible. The delay in cropping caused by variability is mostly due to light exposure and
heavy pruning has an adverse effect on one of the main goals of pruning is to create
early profitability. In intensive plantings the the optimum balance of vigour and fruiting in
vigorous growth caused by heavy pruning as much of the canopy as possible by improv-
may negate much of the advantage in early ing canopy light distribution. The orientation
yields that are achieved by using increased of fruiting branches affects vigour and relative
tree numbers. response to pruning (Fig. 14.5). Normally
upright branches are mostly vegetative and
respond to pruning by growing vigorously. It
14.5.2 Fruiting trees is best to remove these by thinning cuts to
avoid shading portions of the canopy below
Although young trees are often handled them. Pendant branches are often weak and
similarly, as the tree begins to fruit, pruning have low fruit set and poor fruit size, colour
practices must be adjusted to the vigour of and quality (Tustin et al., 1988). These branches
the cultivar and rootstock (see Chapter 5) should be removed to eliminate the small,
and the growth and fruiting habit of the cul- low-quality fruit they produce. Branches with
tivar. Tree vigour is influenced by many fac- an orientation between horizontal and 45º
tors. Fertile, well-drained soil results in above horizontal are the most fruitful, have
much greater vigour than shallow, poorly the highest fruit quality and should form the
drained soil. Some climates are ideal and greatest portion of the fruiting canopy. Even
promote vigorous growth, while others distribution of light across these branches is
cause stresses that reduce growth. Cultural maintained by thinning-out cuts.
practices, such as nitrogen fertilization and Cultivars respond differently to pruning
weed and soil management, influence and training due to differences in their overall
growth. All these factors interact with the level of vigour, growth and fruiting
genetic potential of the scion and rootstock (Table 14.2). Growth habit refers to the overall
and must be considered when judging the growth pattern of the tree, including the
responses of the fruiting tree to pruning. degree of branching, the branch orientation
Fig. 14.5. Influence of branch orientation on growth and fruiting of apple.

Table 14.2. Relative vigour of some commercially important apple cultivars and their growth and fruiting
habit according to Lespinasse (1980).
Low vigour Medium vigour High vigour
‘Braeburn’ (II) ‘Cox’s Orange Pippin’ (III) ‘Cortland’ (IV)

‘Idared’ (II) ‘Delicious’ (II) ‘Granny Smith’ (IV)
‘Jonathan’ (III) ‘Elstar’ (II–III) ‘Gravenstein’
‘Rome Beauty’ (IV) ‘Empire’ (I) ‘McIntosh’ (III)
‘Spur Delicious’ (I) ‘Fuji’ (IV) ‘Mutsu’ (III)
‘Spur Golden Delicious’ (I) ‘Gala’ (III) ‘Northern Spy’ (II)
‘Spur McIntosh’ (I) ‘Golden Delicious’ (III) ‘Spartan’ (II)
‘Tydeman’s Red’ (IV) ‘Jonagold’ (II)
(upright or spreading) and the branch crotch Type II is characterized by standard-habit

angle. Fruiting habit refers to the overall pat- ‘Delicious’. Branching is more frequent than
tern of fruiting and includes fruit position on in Type I and there is a tendency for the
the ends of long or short shoots, age of spurs fruiting zone to move away from the trunk.
producing most of the crop and location of the The trees have a tendency to develop narrow
crop on the scaffold limbs. Lespinasse (1980) crotch angles and benefit from spreading.
developed a system of classification for Type II trees may develop many medium-
growth habits (Fig. 14.6) and these have strong sized branches, resulting in a too dense
implications for the type of pruning required. canopy if not thinned out. No more than
Spur types, characterized by spur strains four permanent branches per tier should be
of ‘Delicious’, are classified as Type I. These retained. More thinning cuts into younger
tend to form few laterals on main scaffold wood are required to induce spurs and retain
limbs and have strong basitonic characteris- the fruiting zone in the tree interior.
tics (the most vigorous growth is at the base Type III cultivars are characterized by
of the tree). Thus the pattern of growth in ‘Golden Delicious’ and tend to be spreading
type I cultivars is characterized by strong with wide crotch angles and frequent
scaffold limbs breaking at the base of the branching. Cultivars of this type tend to bear
tree. The dominance of the central leader is early, with most of the fruit on spurs or
lost quickly unless shoots intended as scaf- terminals of short shoots borne on 2- to
folds are mechanically spread to weaken 4-year-old wood. The fruiting zone on
their growth relative to that of the leader. cultivars in this classification tends to move
Heading cuts may be needed to develop lat- away from the trunk to the outside of the
eral branches on primary scaffolds, which canopy. This requires thinning on the canopy
forces would-be spurs to grow into vegeta- periphery and heading cuts to lateral
tive extension shoots. Due to the sparse branches on the bases of scaffolds to induce
branching habit more scaffolds may be left, renewal of fruiting wood. Trees of this type
but care must be taken that they are spread are the easiest to adapt to a wide range of
vertically along the leader or the dominance training systems and are generally the easiest
of the leader and its growth will be reduced. to manage in the orchard.
Fruiting occurs on numerous short spurs, Type IV, the tip bearers, are characterized
which are long-lived. The zone of fruiting by ‘Rome Beauty’ and ‘Granny Smith’. They
remains close to the trunk and to the base of tend to have upright main scaffold limbs
the scaffolds as long as sufficient light is with narrow crotches and frequent branch-
maintained in these canopy areas. Type I cul- ing. A weeping terminal habit develops since
tivars tend to be prone to biennial bearing most of the crop is produced on the ends of
and can easily become spur-bound and the previous year’s shoots. The lower half of
senescent, particularly if they are propagated many shoots will be devoid of leaves or fruit
on very dwarfing rootstocks. on young trees, a condition known as ‘blind
Fig. 14.6. Lespinasse (1980) classification of apple-cultivar growth habits.
wood’. Delayed spring pruning can often as ‘Empire’, require special treatment during
induce laterals in this branch region, which the developmental years if they are to be suc-
would normally be bare. This growth habit cessfully grown and managed in training
tends to be the most annual in production and systems that depend on a defined leader.
is often associated with high productivity In a young fruiting orchard, leader man-
and large fruit size. agement and, ultimately, tree height require
Low-vigour cultivar-and-rootstock com- consideration. As planting density intensi-
binations can be pruned harder than high- fies, it is critical that one row of trees does
vigour combinations without upsetting the not shade the adjoining row. Several studies
balance of growth and fruiting. Some root- show that a north/south orientation results
stocks, such as Oregon 1, cause upright in more even canopy light distribution. In
growth, while others, such as M.9 or M.9 model studies (Cain, 1972; Palmer, 1989) it
interstems, cause an open-spreading canopy. has been shown that tree height should not
Rootstocks that promote an open-spreading exceed twice the clear alley width. If a 2.5 m
canopy combined with cultivars with little free alley is needed for spray equipment and
tendency to maintain a central leader, such removal of fruit at harvest, then trees should
not exceed 5 m in height. If this is exceeded, fied on young fruiting trees and the indis-
much of the row will be shaded in the after- criminate cuts from mechanical hedging gen-
noon by neighbouring rows. erally result in uneconomic production.
In-row spacing also has an impact on
leader management. Tustin et al. (1998) have
shown that creating rows with a sawtooth 14.5.3 Old or declining trees
profile from creating a gap between leaders
of adjoining trees results in improved light As trees age, fruit size generally declines and
distribution and improved fruit quality. To it requires a more detailed and time-consum-
achieve these goals there must be strong ing pruning job to achieve adequate fruit
vigour in the leader when the trees are size and quality. Trees can become spur-
young. This is achieved by removing shoots bound and perform as old trees even though
competing with the leader. In cultivars such they are not old (Ferree and Forshey, 1988).
as ‘Empire’ that fail to develop strong lead- These trees often have very large spur com-
ers, the leader should be headed by a third plexes with many weak growing points and
each year and then singled to one shoot early have very little shoot growth in the lower
in the growing season. When the tree reaches portion of the canopy. Spur pruning that
the desired height, the leader can be cut back removes 30% of the growing points on each
to an upward-growing shoot each year, just spur complex, plus a series of heading cuts
below the desired height, and the competing over the canopy can improve fruit size in
shoots removed. In supported systems it is these trees.
possible to let the leader fruit and bend over Another problem common in older
and after fruiting cut to one upright shoot orchards is that too many limbs have been
and repeat this process every couple of allowed to develop in the top of the trees.
years. Handling tree height in systems such The upper limbs in the tree shade the lower
as the slender spindle, which requires a limbs, and this uneven light distribution lim-
weak leader, or systems that have more than its production and fruit size in the bottom
one leader is covered in Chapter 15. half of the canopy. Often the productivity
Some very large orchards attempt to and profitability of old trees are limited by
reduce the cost of pruning by mechanically being both top-heavy and spur-bound.
hedging or pruning only every second or A comparison of several corrective prun-
third year. Mechanical pruning unfortu- ing strategies for top-heavy, spur-bound
nately causes mostly heading cuts, and the ‘Starkrimson Delicious’ trees showed that
vigorous upright shoot growth that follows removing several large limbs in the top of
quickly produces heavy shade in the tree, the canopy or making numerous heading
which reduces flower formation and fruit cuts improved light penetration, improved
quality. This effect is particularly bad if fol- fruit size and increased fruit quality, but the
low-up manual pruning is not used to thin reduction in yield in these treatments
out the tree periphery and allow light pene- resulted in no economic benefit (Ferree et al.,
tration to the canopy interior. Even with 1990). Spur pruning produced the smallest
annual follow-up hand-pruning, mechanical increase in fruit size, but also reduced yield
hedging, although successful in maintaining the least, resulting in the greatest cumulative
tree size, has not been economic in maintain- yield and economic value. The outcome of
ing yield and fruit quality (Ferree, 1976b; this study was due in large part to the
Ferree and Lakso, 1979; Ferree and Rhodus, amount and severity of cuts that had to be
1993). In a comparison of pruning six culti- made to achieve the desired canopy shape.
vars annually with limb spreading compared Converting older trees to canopy shapes
with every second or third year without that improve light distribution, such as the
spreading, it was found that the annual treat- palmette leader (Lakso et al., 1989), can be
ment gave the highest absolute return per done with less loss of yield when starting
tree (Ferree and Lakso, 1979). Thus, although with a well-structured central-leader tree
pruning is a costly practice, the cost is justi- (Warrington et al., 1995).
If trees have been neglected and not effects of summer pruning on apple are not
pruned for several years, it is best to make controlled by carbohydrate levels or by the
several large thinning cuts, removing limbs type of carbohydrate present (Taylor and
to open the canopy. It is critical not to prune Ferree, 1986). A redistribution of carbohy-
too excessively in the first year. A few large drates does occur in response to summer
cuts will allow adequate light penetration to pruning and is probably controlled by the
stimulate fruiting wood lower down in the removal of the buds and/or growing points
canopy. In the second and third year, a grad- (Saure, 1987).
ual thinning of the canopy should be done, Quinlan and Preston (1971) showed that
with the focus on renewing the bearing sur- repeated pinching of a growing bourse shoot
face by favouring the retention of young modified the distribution pattern of assimi-
limbs over old ones. If trees have been lates. Movement of photosynthates out of
neglected for more than a couple of years, it the primary leaf towards the growing shoot
is generally not economical to return them to tip was shifted by the pinching to mainly
commercial production. Thus, an economic basipetal transport toward the flower clus-
comparison must be made before deciding ters and the fruitlets, which is the normal
whether to prune or to remove old or direction of transport after terminal bud for-
neglected trees. mation. A similar shift in assimilate distribu-
tion may be responsible for increased fruit
set and improved spur quality in the canopy
14.6 Summer Pruning interior in response to early-summer pruning.
Although summer pruning has been used

for several centuries in some European 14.6.2 Effects on fruits
orchards, the practice was never widely
adapted until more recent times. Saure (1987) Summer pruning has been used primarily to
published a comprehensive review of the improve red fruit colour and fruit quality,
many studies evaluating the impact of the while attempts to control growth by summer
practice on the growth and fruiting of apple. pruning have been less successful. Many
studies show a dramatic improvement in red
fruit colour of difficult-to-colour cultivars,
14.6.1 Effects on physiological processes such as ‘McIntosh’ and ‘Gala’ (Lord and
Greene, 1982; Warrington et al., 1984); how-
Summer pruning affects several of the basic ever, the red colour of cultivars that have a
physiological processes of apple trees (Ferree high colour factor, such as the red strains of
et al., 1984). Photosynthetic rates of basal ‘Delicious’, is often not affected. Summer
leaves remaining after trees were summer- pruning often improves storage quality by
pruned were 11–39% higher than corre- reducing bitter pit, internal breakdown and
sponding leaves on unpruned trees. These water-core (Preston and Perrin, 1974; Myers
rates were equivalent to those in much and Ferree, 1983a). The reduction in these
younger leaves on unpruned trees. As sum- storage disorders is probably related to the
mer-pruning severity increased, the delay in increase in fruit calcium that results from
decline in photosynthesis increased. This summer pruning (Table 14.3). Fruit calcium
delay in the decline of leaf performance was is enhanced by summer pruning early, before
similar to the response induced by increased terminal bud formation and during the time
cytokinin levels. Since the root system of the of active cell division in the fruit, while
summer-pruned trees was not immediately pruning after terminal bud formation has lit-
affected, it might be expected that the root tle effect on fruit calcium concentrations. The
system would continue to produce the same increase in fruit calcium resulting from sum-
quantities of cytokinins as it did prior to mer pruning can often overcome the adverse
pruning, thus increasing the supply to the effect on fruit calcium induced by reduction
remaining leaves. Evidence indicates that the of crop due to frost or excessive growth from
Apples - Chap 14
332
11/4/03
11:01 am
Table 14.3. Effect of summer pruning for 3 years on growth and fruiting of ‘Jonathan’ apple (from Taylor and Ferree, 1984).
Page 332
D.C. Ferree and J.R. Schupp
Marketable fruit (%)
Trunk Canopy
diameter (cm)
area openness1 Firmness Colour Water-
(cm2) (% sky) > 8.0 8.0–7.3 7.3–5.7 (kg) rating2 core3
Control 168a 14b 3.8b 28.8b 65.1a 6.86b 3.8b 0.38b

Summer-pruned 148b 28a 12.7a 48.9a 37.9b 6.98a 4.0a 0.54a
1 Measured by fish-eye lens at tree base.
2 Colour rating: 1 = green to 5 = full red.
3 Water-core rating: 0 = no water-core to 5 = severe water-core throughout cortex.
Values with letters uncommon are statistically different (P = 0.05).

dormant pruning, but the increase is not ade- vigorous trees that exceed their allotted
quate to overcome a general calcium defi- space. Generally, summer pruning is con-
ciency. If too many shoot leaves are removed ducted on the periphery of the tree remov-
in summer pruning, fruit size can be ing vigorous growth by cutting to the
reduced, as well as fruit soluble-solids con- bud-scale scars or to the first spur on 2-year
centration. Modest summer pruning can wood. Growth produced the year following
achieve the increase in fruit colour without summer pruning has been equal to growth
adverse effects on fruit size and soluble from dormant pruned trees (Myers and
solids; thus, these factors must be closely Ferree, 1983b). Marini and Barden (1982)
monitored to avoid negative effects. found no difference in growth, if vigorous
Although early reports indicated that flow- trees were pruned identically, either while
ering and fruit set were enhanced by summer dormant in winter or while growing in sum-
pruning (Saure, 1987), most recent work mer. Forshey et al. (1992) indicate that mod-
shows either no effect or reductions. Heavy erate summer pruning may have little
summer pruning occasionally causes some practical benefit for vegetative vigour, even
buds to break into growth and flower late in if repeated over several years. They indicate
the season, so that bloom and ripening fruit that summer pruning sufficient to reduce
exist at the same time on trees. This is more vegetative vigour is usually accompanied by
prevalent on some cultivars, such as reduced yield, fruit size and/or fruit
‘McIntosh’ and ‘Jonathan’, and the inflores- soluble-solids concentration and may also
cence is often extended rather than compact. produce significant negative side-effects in
Likewise, the influence of summer pruning flowering and winter-hardiness. Thus, any
on fruit set and fruit abscission is unclear, benefit of summer pruning in controlling
with studies reporting contradictory results. growth is temporary and must be coupled
Since all of these factors influence yield, it is with other techniques and management
not surprising that the reports of the influ- practices to achieve a desirable balance of
ence of summer pruning on yield are variable. growth and fruiting.
14.6.3 Effects on growth 14.6.4 Practical implications
Results on container-grown apple trees indi- In reviewing the research on summer prun-
cate that shoot growth, leaf area, stem diam- ing, several principles evolve that have prac-
eter and root growth are reduced by tical significance (Table 14.4). Pruning early,
summer pruning (Plate 14.2; Taylor and before or just after terminal buds form, can
Ferree, 1981). Summer pruning has been increase current-year fruit set by reducing
advocated as a method of devitalizing very competition from rapidly growing shoots.
Table 14.4. Summary of the influence of summer pruning on the growth and fruiting of apple trees.
Characteristic Increased Decreased No change
Leaf photosynthesis X
Leaf carbohydrates X
Fruit calcium X
Fruit colour X
Bitter pit X?
Water-core X?
Fruit size X?
Fruit soluble solids X?
Late-season flowering X
Growth following season X
Pruning at this time will also result in vege- 20th century, many orchards were planted
tative regrowth in close proximity to the cut at a spacing unsuitable for the rootstock/
and the earlier in the season the pruning is scion/training system combination. As a
done, the greater will be the vigour of the result, overcrowding, decreased production
regrowth. Regrowth in the current season is and lower fruit quality occurred. A means
reduced or eliminated if summer pruning is was needed to control tree growth, and root
performed 1–2 months before harvest but pruning was one of the techniques evalu-
after terminal buds have formed. Regrowth ated. A review of root pruning (Geisler and
in the current season is also reduced as cuts Ferree, 1984b) reported that it was widely
are made into older wood. Cuts made to the practised in European gardens to reduce tree
first spur bearing fruit on 2- or 3-year-old size and promote flowering in the 1800s. In
wood often do the most to open up the later years it was considered too dwarfing to
canopy and promote improved fruit colour, be a recommended practice in either
without undesirable current-season re- European or North American orchards.
growth. Removal of entire shoots by thin-
ning cuts produces little or no regrowth
when done after terminal-bud formation. 14.7.1 Effects on physiological processes
Because the remaining sites of auxin pro-
duction are undisturbed, little disruption of Root pruning of non-bearing apple trees in
the apical-dominance control system occurs. the greenhouse temporarily reduces net
This approach provides the maximum photosynthesis, transpiration and leaf water
improvement in fruit colour since it allows potential (Geisler and Ferree, 1984b; Schupp
more light to reach the fruit. and Ferree, 1990). When roots are removed
Although the primary use of summer by pruning, assimilates are redistributed to
pruning is on fruiting trees to improve fruit the root system, as evidenced by the
colour and quality, there are several other increase in small roots in close proximity to
uses that are beneficial. Removal of water the trunk, coupled with a reduction in leaf
sprouts in the tree centres and on major scaf- and new shoot growth. In a field study, root-
folds of older trees improves light and spray pruned trees had more negative leaf water
penetration and reduces problems from potential, reduced stomatal conductance
some insects. Water-sprout removal can be and reduced transpiration (Schupp et al.,
done any time, but pulling them off early in 1992). There was little effect on concentra-
the season is preferred since cutting later tions of leaf nutrient elements (Schupp and
after the wood hardens leaves basal buds to Ferree, 1987; Schupp et al., 1992).
regrow. Summer pruning can be used on Shoot-tip ethylene production was
young trees to encourage leader develop- reduced at 3, 9 and 17 days after root prun-
ment in such systems as the central leader, ing, while the concentration of the ethylene
slender spindle and central axis (see precursor, 1-aminocyclopropane-1-carboxylic
Chapter 15). Normally it is best to remove acid, in the xylem sap was unaffected. The
several newly developing shoots just below concentration of cytokinin in the xylem sap
the leader as soon as growth is 2–5 cm long. was reduced 1, 3 and 9 days after root prun-
Summer pruning should not be used on ing (Schupp and Ferree, 1994). Thus, ethylene
trees with low vigour. The loss of leaf sur- and cytokinin did not appear to be responsi-
face and the photosynthetic capability it rep- ble for the reductions in growth caused by
resents can only aggravate the pre-existing root pruning.
vigour problem. Although root pruning reduces photosyn-
thesis and transpiration, it has no influence
on the carbohydrate fractions in the leaves or
14.7 Root Pruning shoots, but causes an increase in soluble and
insoluble fractions in the roots (Ferree, 1989).
As orchard intensification became a world- The consistent water-stress effect found from
wide phenomenon in the latter half of the root pruning probably explains most of the
growth response, as growth rate recovers Compared with young greenhouse-grown

along with regeneration of the root system trees, the effects of root pruning are much
and its ability to meet evapotranspirational greater and of longer duration on bearing
demand. During the interim stress, the tree trees in the orchard. The presence of crop
reduces water use by stomatal closure and and its effect on root regeneration explain
minimizes transpirational surface through the stronger response. For example, the
reduced shoot and leaf growth. The shift in annual increase in trunk cross-sectional area
plant hormones may also play a role directly of fruiting trees was reduced 44% by root
by influencing growth or indirectly by influ- pruning, but only 14% in deblossomed trees
encing water relations. (Schupp et al., 1992). The presence of crop
reduced root regeneration and was, there-
fore, of primary importance for season-long
14.7.2 Effects on growth reduction in vegetative control by root prun-
ing. The resulting long-term water deficit
Results from greenhouse studies on young is thought to be the means by which shoot
apple trees showed that root pruning growth is reduced. Lenz (1986) showed
decreased leaf number per tree, leaf size, that cropping trees increased water usage
total leaf area, shoot growth and decreased compared with non-cropping trees. In a
dry weight of leaves, shoots and roots 6-year root-pruning study, shoot growth of
(Geisler and Ferree, 1984a; Ferree, 1989; ‘Jonathan’ apple was reduced approximately
Schupp and Ferree, 1990). In the field, root 30% each year (Fig. 14.7a), except the single
pruning reduced trunk cross-sectional area, year where rainfall each month of the grow-
shoot length, shoot number to spur ratio, ing season greatly exceeded the long-term
shoot leaf size, and shoot leaf area to spur- average (Ferree, 1992).
leaf area in cropping apple trees (Schupp Root pruning results in root regeneration
and Ferree, 1987, 1988; Ferree, 1992). in close proximity to the cuts. The timing of
The overall growth effects resulting from root pruning affects how much root regener-
root pruning were sufficient to reduce prun- ation takes place in a season. Considerable
ing time and increase within-canopy light root regeneration was evident at the end of
levels. Canopy light penetration and spur the growing season on trees root-pruned
quality were increased. The more severe (the while dormant or at full bloom. Root regen-
closer to the trunk) the root pruning, the eration was less on roots pruned at June
greater were the effects on growth. drop and minimal in preharvest root-pruned
(a) (b)
40 125
a a a
Average shoot length (cm)
35 a a a
30
Yield (kg per tree)
a
100 b
25 a
b b b
20 b
b a a
15 b b
75
10 b
Control b Control
5 Root pruned (bloom) Root pruned (bloom)
0 50
85 86 87 88 89 90 85 86 87 88 89 90
Year Year
Fig. 14.7. Influence of root pruning over 6 years on shoot growth (a) and yield (b) of mature ‘Jonathan’
apple trees. Note the consistent growth effect and reduction in biennial bearing. (From Ferree, 1992.)
trees (Schupp and Ferree, 1987). Excavations 50 cm, there was no difference in response,
around trees root-pruned annually for 9 as the upper 25 cm of soil has the greater
years showed a reduction in all root sizes. concentration of roots. The optimum time of
Roots < 1 mm in diameter were reduced 20% pruning appears to be around full bloom. In
by root pruning, while the reduction in a study comparing responses of nine root-
larger roots was nearly double this amount stocks or interstock combinations to the
(Ferree, 1994a). The effect of root pruning on effects of root pruning, growth on all trees
root distribution was greatest in the top was reduced 20–30%, with no difference
30 cm of soil, parallel to the location of the whether it was on dwarfing M.9 or vigorous
root-pruning cut at a depth of 25 cm. Roots seedling rootstock (Ferree and Knee, 1997).
below 30 cm were unaffected. Multiple spray applications of low concen-
trations of cytokinins in the field were not
successful in counteracting the reduction in
14.7.3 Effects on fruiting fruit size due to root pruning.
Since root pruning has the potential to
Compared with unpruned trees, root prun- control tree size, a 10-year study evaluated
ing increases yield efficiency (yield per trunk the potential of root-pruning four cultivars
cross-sectional area), fruit colour and soluble- grown in two orchard systems (Ferree and
solids concentration. Preharvest fruit drop Rhodus, 1993). Root pruning was successful
and fruit size are reduced. In large-fruited in containing tree size of apple trees planted
cultivars (‘Melrose’, ‘Golden Delicious’) the at half the recommended in-row spacing.
effect on fruit size was not enough to reduce However, the reduced tree efficiency in the
yields, but in small-fruited cultivars, such as trellis system of dwarf trees or in the central-
‘Jonathan’, yield was reduced (Fig. 14.7b). leader system of semi-dwarf trees resulted in
Root pruning at full bloom increased minimal cumulative yield increase and no
fruit soluble-solids concentration and apparent economic advantage. It was con-
slightly increased fruit firmness but cluded that it is better to plant systems at a
decreased starch hydrolysis at harvest spacing that avoids excessive crowding and
(Schupp et al., 1992). Root pruning does not allows an efficient balance of growth and
appear to affect blossom density or fruit set. fruiting without the need to intervene with
In a long-term study over 9 years, root prun- practices such as root pruning.
ing reduced the biennial bearing pattern. Root pruning is a mechanical means of
Elfving et al. (1991) found that root pruning tree size and vigour control, which can assist
at full bloom in ‘McIntosh’ reduced the inci- in such situations as improper spacing for
dence of stem-cavity browning and brown rootstock or cultivar or loss of crop to frost
core, but not in every year. Root pruning did or following a particularly hard pruning to
not influence ethylene evolution at harvest, bring growth and fruiting balance to trees
but did reduce post-storage ethylene. They (Table 14.5). Under current market condi-
concluded that root pruning at full bloom tions there is a strong emphasis on produc-
was generally less effective than daminozide ing large-sized fruit to satisfy the
in altering fruit behaviour. preferences of apple retailers. Since root
pruning reduces fruit size, it must be
viewed as a ‘rescue treatment’ for orchards
14.7.4 Practical implications with excessive growth and overcrowding,
rather than a standard practice for reducing
Generally, the closer to the trunk that root growth and pruning labour in orchards with
pruning is carried out, the more roots are cut more normal vigour.
and the greater the effect. For bearing trees, It is important to understand the effects of
cuts 70–100 cm from the trunk on two sides root pruning since all trees purchased from a
are required to have the effects on growth nursery have lost a portion of their root sys-
and fruiting previously described (Plate 14.3). tem. Several reports indicate no effect of
In a study comparing depths of 25 and removing most of the roots from nursery
Table 14.5. Summary of the influence of root pruning on the growth and fruiting of apple trees.
Characteristic Increased Decreased No change
Trunk growth X
Shoot length X
Shoot number X
Spur/shoot ratio X
Shoot-leaf size X
Spur quality X
Pruning time X
Canopy light penetration X
Biennial bearing X
Fruit set X
Fruit yield (no. of fruit) X
Fruit size X
Preharvest drop X
Fruit colour and quality X
Tree-yield efficiency X
trees after they are dug up on subsequent of cells is changed and reaction wood is pro-
tree growth. However, trees propagated duced on the upper side of the branch.
in the field and transplanted under ideal Reaction wood consists of xylem tissue on
conditions caused a 32% reduction in growth one side of the stem that has short, round
compared with trees propagated in the field tracheids and an altered pattern of wall
and not transplanted (Ferree, 1976a). Thus, thickening. Reaction wood can often be
the effect of root pruning can be significant observed on the pruning cut of side limbs,
and care should be taken not to sever roots with the upper half appearing denser than
through cultivation or other cultural prac- the lower half.
tices if no reduction in growth is desired. The stress created by bending has been
shown to result in increased ethylene con-
tent in the internal air spaces within the
14.8 Training branch (Robitaille and Leopold, 1974). The
ethylene reaches a maximum 2 days after
Training utilizes various techniques, includ- bending stress is imposed and returns to
ing pruning, to direct tree growth or form control levels in 3 weeks. A similar increase
and the development of the structural in ethylene was shown to occur by a paste
framework of the tree. Although these forms application of auxin (naphthaleneacetic acid
can be very ornamental (Plate 14.4), gener- (NAA)) to the branch, which in turn caused
ally the tree form fits some predesigned an increase in ethylene. Thus, both auxin
training system described in Chapter 15. and ethylene may play a role in the
Discussion in this chapter will be restricted responses induced by bending.
to the techniques used to control tree growth The growth reduction and increase in
in many orchards. flower initiation caused by bending occur
only during the season when the physical
bending stress is imposed or the limb orien-
14.8.1 Bending tation is changed (Table 14.6). This effect has
commercial value, since, by progressively
Physically bending a branch of an apple tree changing branch orientation by inserting a
results in a reduction in terminal shoot longer spreader each year, a growth reduc-
growth and a redistribution of growth hor- tion can be achieved for several years. The
mones, particularly auxin. The arrangement progressive influence of spreading the stress
Table 14.6. Influence of bending on apple growth and flowering in the year of treatment (from Ferree,
1994b).
Shoot length Limb area Flowers

Treatment (cm) (cm2) per limb area
Control 26.8a 2.14b 13.2b

Bending 24.4b 2.48a 15.3a
Values with letters uncommon are statistically different (P = 0.05).
over 2–3 years results in the greatest num- The bending of a branch influences not
ber of short, fruitful shoots and the fewest only the amount of growth that occurs
number of undesirable shoots. Cropping in but also the location where shoots arise
successive years has a similar influence on (Fig. 14.8). The influences of auxin begins
the tree, but it is not as predictable as physi- just below the shoot tip and, as the orienta-
cally bending and does not occur until after tion away from vertical increases, the auxin
the limb may have grown for several years influence progresses down the branch and
and stiffened in an upright position, requir- buds on the upper side are released from the
ing considerable force or weight to change hormonally induced apical dominance and
its orientation. The shoot-growth reduction begin to grow. Thus, the more the branch ori-
that results from bending is associated with entation is changed towards the horizontal,
a similar reduction in root growth, so that the greater the number of shoots released
the top-to-root ratio of the young tree is towards the branch base. If basal shoots are
maintained. released due to bending too severely, they
Forshey et al. (1992) suggested that the tend to be very vigorous and upright and
reduced vegetative vigour in limbs that are often require removal during the dormant
bent reduces the production of gibberellins, season. By bending and changing orientation
which are antagonistic to flowering. The for- only 10–15° in any year, shorter shoots are
mation of moderately vigorous lateral shoots produced mostly on the outer half of the
and spurs creates additional sites for flower branch. If excessive spreading or a heavy
formation. These two effects result in crop load on the tip of the branch brings the
increased flowering and earlier fruiting. branch to a horizontal or below-horizontal
Fruit quality is also improved because of position, vegetative growth on the weeping
improved light penetration and because tip portion nearly ceases and several very
more fruit hang free instead of rubbing vigorous suckers will form from buds on the
against the branch on more upright limbs. highest point of the bend. This undesirable
40° 60° 70° 90°
Fig. 14.8. Influence of various degrees of bending on distribution and amount of growth of an apple branch.
type of response can be avoided by not always found. In training systems such as
bending the limb beyond about 60° from ver- the slender spindle or hybrid tree cone
tical and avoiding overcropping on the tips orchard system (HYTEC), bending is used
of scaffolds of young trees. to weaken the leader and encourage
Several reports indicate that the inclina- growth and fruiting in the lower scaffolds.
tion of the scion, particularly if horizontal, In a study that compared the influence of
reduced growth and increased flowering bending the leader first one way and then
(Tromp, 1972). The study concluded that the the opposite way for different lengths of
reduction of growth and promotion of flow- time, the length of time the leader was bent
ering are independent phenomena and shoot had no effect on shoot growth, but 15 days
orientation interferes in the later phases of appeared to increase flowering, with no
flower formation. The effect on flowering benefit from longer times in the bent posi-
was equivalent to that caused by damino- tion (Ferree and Schmid, 1999). The
zide. A field study over 5 years with ‘Golden hypothesis that the response to bending
Delicious’/M.9 found that inclining the may differ at various times during spring
trunk at planting 45° or 60° from the vertical when growth is initiated was refuted in this
reduced the rate of shoot elongation by 40% study, as no effect was found (Fig. 14.9).
and trunk circumference growth by 17% and Bending in combination with minimal
27%, respectively (Bargioni et al., 1995). Root pruning was successful in reducing growth
total dry weight was reduced by inclination and increasing production of young ‘Fuji’
and roots were more numerous and more trees (Ferree and Schmid, 1994). In a study
elongated in the direction of inclination. The with two cultivars that compared responses
authors propose that the physiological basis on the dwarfing rootstock Mark and the
of the phenomenon may reside in the asym- semi-dwarf rootstock M.7a, bending scaf-
metric distribution of hormones caused by folds increased flowering in the 2 subse-
the inclination. These effects of inclination quent years and increased cumulative yield
probably influence trees trained in such 11% on Mark, but had no effect on the yield
systems as the Lincoln canopy or Tatura of trees on M.7 (Ferree, 1994b). It is unclear
trellis, where trunks or branches are inclined why bending provides an economic benefit
and secured to the trellis framework. by reducing or redirecting growth in some
Practically, bending is accomplished using instances and in others the effects are negli-
many techniques. In central-leader training, gible.
scaffold limbs are spread using wooden,
metal or plastic spreaders, either notched or
pointed to keep them in place. These are nor- 14.8.2 Phloem interruption
mally inserted before growth starts and can
be removed after 4–6 weeks of growth. Others Scoring and ringing, which cause the inter-
prefer to bend by tying limbs down to other ruption of the downward flow of carbohy-
limbs or anchors in the soil or fastening to the drates and hormones in the phloem, have
support system. The ties must be watched been shown to alter growth and fruiting in
and cut if they begin to girdle the limb. apple. The degree or length of time that
Weights have also been used and again care translocation through the phloem is inter-
must be given to avoid girdling. Spreaders rupted is influenced by whether and how
and ties move a limb to a given position, but much of the bark and phloem are removed.
with weights it is more difficult to know Scoring, which is a circumscribing cut
exactly the degree of bending that will occur. through the bark, but not into the wood, has
Special rubber bands can be used to bend the smallest effect. Ringing, which removes
actively growing shoots or a leader, and again different amounts of bark in a circumscribing
the degree of bending is difficult to ascertain. ring, increases in severity with the amount of
Although bending has been commonly bark that is removed until the ultimate of
used to reduce shoot extension and pro- tree death occurs when the tree is unable to
mote flowering, consistent effects were not form new tissue to bridge the ring.
70
LSD = 11.5
60
Terminal shoot length (cm) 50
40
30
Check
20
Bend
10 Head
Bend + head
0
16 Apr. 25 Apr. 9 May 23 May 6 June 20 June
Time of bending
Fig. 14.9. Interaction between time of bending and heading vigorous laterals on the central leader of ‘Regal
Gala’ on M.7 rootstock (from Ferree and Schmid, 1999).
Scoring and/or ringing interrupt phloem Elfving et al. (1991) found that trunk scoring or
translocation, which causes a build-up of the ringing increased soluble solids and retarded
products of photosynthesis in the portion of loss of flesh firmness before harvest and fol-
the tree above the cut. Growth hormones pro- lowing storage, but had little effect on starch
duced in the apical meristem and young hydrolysis. Scoring or ringing decreased the
leaves are also translocated in the phloem and incidence of brown core, stem-cavity brown-
thus also play a role in the effects produced. ing and scald in some years. Scoring also
The photosynthetic and transpiration rates of reduced post-storage ethylene evolution.
leaves above the scoring cut are reduced for ‘McIntosh’ fruit were longer and had larger
up to 2 weeks, but recover to the control lev- pedicel diameters and weights and more
els of unscored plants in 4 weeks. This effect colour from trees that were ringed (Webster
is thought to be due to the carbohydrate feed- and Crowe, 1969). Results from these studies
back mechanism, which causes photosynthe- were variable on the effects of phloem inter-
sis to slow or stop when carbohydrates build ruption on fruit size and quality and were
up and are not translocated out of the leaves. often not consistent from year to year.
If a ring of bark is removed, the effects on More severe versions of phloem interrup-
photosynthesis last for more than 4 weeks tion (Plate 14.5) can be done with a chainsaw
and a greater reduction in growth would be (Hoying and Robinson, 1992). Chainsaw ring
expected. A number of studies show that girdling involves making two overlapping
phloem interruption reduced terminal horizontal rings, 2 cm deep and spaced 20 cm
growth, trunk cross-sectional area, shoot apart on the trunk. With guillotine girdling,
diameter and number of shoots (Greene and two chainsaw cuts were made to remove all
Lord, 1983; Autio and Greene, 1992). the tissue on opposite sides of the trunk to a
Trunk scoring 19 days after full bloom depth of one-third the radius of the trunk.
reduced fruit size in ‘Cortland’ and ‘Delicious’ Both techniques reduced growth, although not
apples, reduced firmness in ‘Delicious’ and as much as root pruning, with the advantage
increased soluble solids in both cultivars that fruit size was not reduced by girdling
(Greene and Lord, 1978). After storage, (Hoying and Robinson, 1992). Caution must
‘Cortland’ apples from scored trees had a be exercised when applying herbicides under
higher incidence of bitter pit and cork, while trees that have undergone these treatments to
‘Delicious’ fruit exhibited more breakdown. avoid damaging the callus tissue that forms as
the girdles heal. Furthermore, this treatment the bud, but it was not able to overcome inhi-
has induced the development of heart-rot in bition due to geotropism. Shoots that develop
some trees. It is cautioned that this technique from notched buds usually have a sharper
should be used only to extend the economic crotch angle than shoots that develop from
life of orchards that are nearing the end of buds that were not notched (Verner, 1955).
their useful lifespan (Hoying, 1993). Crotch angle can be improved by using
clothes-pegs or other techniques, as previ-
ously described. Greene and Autio (1994)
14.8.3 Notching found notching consistently effective, indicat-
ing that one could reasonably expect more
Notching is a phloem-interruption technique than 80% of the notched buds to grow into
used to stimulate lateral branching from buds lateral shoots. Notching is a very useful tech-
that would not normally grow. The technique nique for inducing shoots for scaffolds and
removes a 2–3 mm wide strip of bark directly for balancing growth in young trees. In addi-
above a bud. The cut extends down to the tion to using pruning, bending and notching
secondary xylem and around about one-third to induce shoot growth, chemical growth reg-
of the circumference of the stem. Notching ulators (Chapter 17) and the removal of the
stimulates lateral branching by interrupting youngest cupped leaves on the apical meris-
the downward movement of auxin from the tem (Chapter 6) can be used in appropriate
shoot tip (Tamas, 1987). As long as the situations. Enclosing the base of very vigor-
phloem remains severed and auxin is pre- ous upright leaders in plastic sleeves just
vented from reaching the lateral bud, the prior to the beginning of growth has been
growth and development of the bud will be used to stimulate buds to grow (Plate 14.6).
induced. Greene and Autio (1994) found that
the most efficient time to notch was approxi-
mately 2–4 weeks before full bloom. Buds 14.9 Conclusion
notched shortly after bloom grew less than
those notched earlier. Although notching Pruning and training occupy a unique posi-
increased the growth of all bud sizes, the tion in the management of an orchard, being
largest buds were more likely to develop into the most costly cultural practices and the
a lateral shoot and the shoot would be longer practices most often poorly done, with dra-
than from small buds. Notching was most matic influences on orchard efficiency and
effective at inducing shoot growth from buds fruit quality. Pruning interacts with other
on the top of the branch, less effective for cultural practices, such as fertilization, fruit
buds on the side and least effective for buds thinning, growth regulators and pest control.
on the underside of the branch. The percent- Fruit quality is a sensitive indicator of the
age of buds that developed into shoots was effectiveness of pruning, as canopy light dis-
not influenced by wood age or location along tribution influences all aspects of yield and
the length of the shoot. Notching was able to fruit quality. Although good pruning does
overcome bud inhibition imposed by a shoot not guarantee success, economic success
apex by preventing the movement of auxin to without good pruning is inconceivable.
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Myers, S.C. and Ferree, D.C. (1983a) Influence of time of summer pruning and limb orientation on yield,
fruit size, and quality of vigorous ‘Delicious’ apple trees. Journal of the American Society for
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15 Apple-orchard Planting Systems
Terence L. Robinson
Department of Horticultural Sciences, New York State Agricultural Experiment Station,
Cornell University, Geneva, New York, USA

15.2 Spherical-shaped Canopy Systems 347
15.2.1 The bush-tree system 347
15.2.2 Spindle-bush system 347
15.2.3 Advantages and disadvantages of spherical-shaped systems 347
15.3 Conic-shaped Canopy Systems 348
15.3.1 Central-leader system 348
15.3.2 Mini-central-leader system 348
15.3.3 Palmette-leader system 350
15.3.4 Slender-spindle system 351
15.3.5 North Holland spindle 353
15.3.6 Slender-spindle multi-row or bed system 354
15.3.7 Vertical-axis system 354
15.3.8 SolAxe system 356
15.3.9 Slender-pyramid system 357
15.3.10 HYTEC system 358
15.3.11 Super-spindle system 362
15.3.12 Meadow-orchard system 363
15.3.13 Advantages and disadvantages of conic-shaped systems 363
15.4 Flat Planar Canopy Systems 365
15.4.1 ‘Regular’ palmette system 365
15.4.2 ‘Free’ palmette system 366
15.4.3 Penn State thin-wall trellis system 368
15.4.4 Lincoln canopy system 369
15.4.5 Ebro trellis system 369
15.4.6 Solen system 370
15.4.7 Tabletop bed system 371
15.4.8 Present status of flat planar systems 371
15.5 V-shaped Canopy Systems 371
15.5.1 Tatura trellis system 372
15.5.2 Mini-Tatura trellis system 372
15.5.3 Geneva Y-trellis system 372
15.5.4 Mikado and Drilling systems 373
15.5.5 MIA trellis system (A-shaped trellis) 373
15.5.6 Mini-V-trellis system 373
15.5.7 Gütingen V slender-spindle system 375
15.5.8 V super-spindle system 375
15.5.9 Advantages and disadvantages of V systems 377
346 T.L. Robinson
15.6 Tree-support Systems 378

15.7 Field Comparisons of Orchard Planting Systems 380
15.8 Underlying Principles of High-density Orchard Planting Systems 382
15.8.1 Tree density 382
15.8.2 Rootstock 384
15.8.3 High light interception at orchard maturity 385
15.8.4 Good light distribution within the mature canopy 387
15.8.5 Balance between vegetative growth and cropping 392
15.9 Orchard Management Practices for Successful High-density Orchards 393
15.9.1 Soil and site selection 393
15.9.2 Tree quality 394
15.9.3 Time of planting 394
15.9.4 Weed control 394
15.9.5 Irrigation 394
15.9.6 Fertilization 395
15.9.7 Soil fumigation 395
15.10 Economic Comparisons of Orchard Planting Systems 395
15.1 Introduction considerable variability in grower success

with each planting system (Hoying and
Orchard planting systems are specific com- Robinson, 2000).
binations of orchard layout and manage- Modern orchard planting systems are
ment that are designed to improve orchard based on higher tree densities than were
production efficiency. They include the common 50 years ago when most orchards
management variables of rootstock, tree were planted at a density of 70–100 trees
spacing, tree arrangement, canopy shape, ha1. Today tree densities of modern apple
tree-training method, pruning method and orchards range from 1000 to 6000 trees ha1,
tree support system. Over the last 30–40 with some systems using densities up to
years, numerous planting systems for mod- 10,000 trees ha1. This increase in tree
ern apple orchards have been developed, density has been made possible by the
each with their own merits. Although the development of dwarfing apple rootstocks.
systems differ in specific management The combination of dwarfing rootstocks and
practices, many of the modern planting higher tree-planting densities has dramati-
systems have similarities and are based on cally improved cumulative fruit production
the same underlying principles. They all over the first 10 years of an orchard’s life.
have the goals of high early yields, high With most modern high-density planting
sustained yields and excellent fruit quality. systems, a small but significant yield is
Growers can achieve these goals with any expected during the second growing season
of several orchard planting systems. There of the orchard. Substantial yields are
may be no perfect planting system for all expected in the third year and mature yields
growers, but how a fruit grower integrates are expected by year 5 or 6. In contrast, tra-
the practical management variables of each ditional low-density systems on vigorous
system determines his/her opportunity for rootstocks began production around year 6
economic success with the new orchard. or 7 and did not reach mature yields until
Barritt (1992) has likened each of the man- year 15.
agement variables that make up an orchard Orchard planting systems can be catego-
planting system to puzzle pieces. Fruit rized by tree canopy shape. There are four
growers must successfully integrate these basic shapes of canopies: spherical canopy
puzzle pieces to be profitable. Ignoring one shapes, conic canopy shapes, flat-fan
or more of the puzzle pieces has resulted in canopy shapes and Y or V canopy shapes.
Apple-orchard Planting Systems 347
European gardeners have utilized various 15.2.1 The bush-tree system

clonal dwarfing and semi-dwarfing root-
stocks for centuries and have trained apple In the 1930s, as research results from clonal
canopies into many geometric shapes dwarfing rootstock trials were obtained, a
(Huggard, 1980). The most common of smaller version of the large globe-shaped
these were the espaliers grown along walls tree on clonal rootstocks, named the bush
and fences. However, most commercial tree, was developed in Holland and
apple orchards in the 1800s and early 1900s England (Wertheim, 1981). The tree canopy
utilized trees on seedling rootstocks and was lowered to only 50 cm above the
had large globular-shaped tree canopies. ground, which no longer allowed animal
Other tree shapes were not an integral part grazing in apple orchards. The tree height
of commercial orchard planting systems was reduced from 6–8 m to 4–5 m and tree
until the middle of the 20th century. In the density was increased from 70–100 trees
second half of the 20th century, researchers ha1 to 250–350 trees ha1. This tree system
and growers began to search for more effi- had earlier production, due to the use of
cient shapes and systems. There have been semi-dwarfing rootstocks, easier manage-
nine international symposia in the last 40 ment, due to the smaller trees, and higher
years convened by the International Society orchard productivity.
for Horticultural Science’s working group
on Orchard and Plantation Systems that
have chronicled the developments in high- 15.2.2 Spindle-bush system
density orchard systems over the last 50
years. This collection of papers, published In the late 1930s the spindle-bush system was
in Acta Horticulturae volumes 65, 114, 160, developed in Germany by Schmitz-Hübsch
243, 322, 349, 451, 513 and 557, is suggested and Heinrichs (1942), which revolutionized
as further reading. apple growing (Wertheim, 1981). The spin-
dle-bush was based on the fully dwarfing
M.9 rootstock and utilized, for that time, high
15.2 Spherical-shaped Canopy Systems tree densities of 1000–1500 trees ha1. The
trees were spaced 2.5 m in the row and had a
The most common tree form in traditional tree height of only 2 m. The tree had a single
apple orchards in Europe and North leader and was supported by a pole. The hor-
America in the late 1800s and early 1900s izontal fruiting branches originated from the
was a large globe-shaped tree with a height leader. The use of M.9 and high tree densities
of 6–8 m and a main trunk length of 1.8–2 m, gave very high early production, easy man-
which provided enough room under the agement and good fruit quality. The use of
canopy for cattle to graze (Walker, 1980). The the spindle-bush system was short-lived, but
tree was planted on a seedling rootstock, it led directly to the slender-spindle system in
using a low tree density that required large the 1960s, which has become one of the lead-
spreading trees to fill the space allocated to ing systems in the world.
each tree. This required multiple large scaf-
fold branches or leaders per tree, which
formed large globe-shaped canopies at matu- 15.2.3 Advantages and disadvantages of
rity and required 10–15 years to produce spherical-shaped systems
high yields (Plate 15.1). In the 1920s, Sir
Ronald Hatton of East Malling Research The primary advantage of spherical shapes
Station collected, categorized and then is that they are generally the natural shape
released the Malling series of clonal dwarf- produced by apple trees. However, the
ing rootstocks from all over Europe, which canopies generally have a large shaded cen-
spurred the commercial development of tre core that is unproductive and produces
orchard systems based on smaller trees poor fruit quality. In addition the traditional
(Walker, 1980). spherical canopies, although large at matu-
348 T.L. Robinson
rity, required too much time to develop. distance between the bottom and second
Globe-shaped canopy systems were the tier to a minimum of 1 m and to leave gaps
dominant tree form through the first half of in the canopy to increase the light penetra-
the 20th century but were replaced by conic, tion to the bottom tier.
planar and V systems in the later half of the The canopy is developed by heading the
century. None of the current leading leader annually to develop tiers of branches
orchard systems is based upon a spherical just below the heading cut (Table 15.1).
canopy shape. Branches in each tier are manually spread to
an angle of 30° above horizontal for the bot-
tom tier and to horizontal for the upper tiers.
15.3 Conic-shaped Canopy Systems Upper-tier branches are shortened when they
become too long to maintain the conic shape.
In the early 1960s, researchers (Heinicke, The New Zealand version of the central-
1963, 1964; Looney, 1968; Cain, 1970, 1972) leader tree maintains gaps between branches
studied the light distribution within the in each tier for ladder bays and light penetra-
canopies of large globe-shaped apple cano- tion to the centre of the tree.
pies and concluded that much of the canopy
received too little light for good fruit quality
and was unproductive. They proposed a 15.3.2 Mini-central-leader system
conic or pyramidal canopy shape as an
improved tree form. This tree form was then The mini-central leader was developed by R.
developed into several planting systems. Norton in New York State in the late 1970s. It
is similar to the central leader, but utilizes
semi-dwarfing rootstocks and slightly higher
15.3.1 Central-leader system tree densities, ranging from 500 to 1000 trees
ha1. Trees are typically supported with an
The central-leader system was developed individual 2.5 m tree stake. Trees are pyramid-
by Heinicke (1975) in North America and shaped, with two tiers of permanent branches
McKenzie (1972) in New Zealand. This along the trunk and a tree height of 3–4 m.
tree-training system has been widely Semi-dwarfing rootstocks, such as M.26,
adopted in North America, New Zealand, M.9/MM.111 or M.9/MM.106 interstems,
Australia and South Africa. The system has G.30 and Supporter 4, are used.
a pyramid-shaped tree with tiers of
branches spaced along the trunk. It is a 4.5 m
medium-density system with tree densities
ranging from 300 to 700 trees ha1.
Typically trees have three to four tiers of
branches spaced along the trunk and a tree
height of 4–5 m (Fig. 15.1 and Plate 15.2).
The trees are usually not supported with a
trellis or individual tree stakes. Semi-vigor-
ous rootstocks, such as M.7, MM.106,
MM.111 and M.793, are used. The widest 1m
part of the tree is at the bottom tier. The
bottom tier of branches is 60–90 cm above
the ground and has four or five branches.
The second tier of branches is commonly
60 cm to 1 m above the first tier and has 3m
four branches. The third and fourth tiers
are spaced about 60–80 cm apart and typi- Fig. 15.1. The central-leader tree has a free-
cally have three branches. Years of experi- standing dominant trunk, a conic shape and distinct
ence have led growers to increase the tiers of branches, with a 1 m gap between tiers.
Table 15.1. Simplified pruning and training plan for the central-leader system. (NB. Northern-hemisphere
dates.) (Adapted from Robinson and Hoying, 1994.)
First year
At planting Adjust graft union to 10 cm above soil level. Remove all scaffolds below 60 cm,
using a flush cut. Trees with three or more scaffolds (25 cm long) should be headed
at 110 cm above the soil line, with all scaffolds headed by removing one-third their
length. Trees with fewer than three feathers should be headed at 90 cm and all
feathers removed, using a bevel cut, leaving a 2 cm stub
1–2 cm growth Rub off second and third buds below the new leader bud to eliminate competitors to
the leader shoot. Deflower tree
5–10 cm growth Attach clothes-pegs to new side-shoots on leader to promote wide crotch
angles
Early summer Install tree-support system that will allow tree to be supported to 3 m. Attach tree to
support system with a permanent tree tie above first tier of scaffolds, leaving a 5 cm
diameter loop to allow for trunk growth
July Tie developing leader to support pole with permanent tie. Remove clothes-pegs
Second year
Dormant Head leader, removing one-third to two-thirds of last year’s growth, depending on
tree vigour. The more vigorous the leader and scaffolds, the less severe the
heading cut should be. Remove any side-branches remaining that compete with the
leader. Scaffold branches should be headed, removing one-third of last year’s
growth
5–10 cm growth Attach clothes-pegs to lateral shoots developing on 1-year-old wood to ensure good
crotch angles. Select leader and pinch out or remove competing shoots
Mid-July Spread bottom-tier branches to 45° using 30 cm nail-tipped limb spreaders.
Spreading should be done early in the growing season for vigorously growing trees,
later for weaker-growing trees. Remove clothes-pegs
Third year
Dormant Head leader 1 m above the top branch of the bottom tier of branches to promote
permanent second-tier scaffolds. Remove vigorous branches between first and
second tiers of branches
5–10 cm growth Attach clothes-pegs to lateral shoots developing on 1-year-old wood to ensure good
crotch angles. These will form permanent second tier. Select leaders and pinch out
or remove competing shoots
August Remove clothes-pegs
Fourth year
Dormant Head the leader, removing one-third to one-half of last year’s growth. Remove
excessively vigorous non-scaffold limbs, particularly those competing with the leader.
Reposition 30 cm spreaders where needed within the tree
July If bottom-scaffold branch length is approaching one-half in-row spacing, respread
with 1 m nail-tipped spreaders to 80° from vertical
August Lightly summer-prune to encourage light penetration and maintain pyramidal tree
shape
Fifth year
Dormant Head leader, removing one-third of last year’s growth. Remove shoots that are
competing with the leader. Lightly dormant-prune to invigorate weak areas, thin out
overcrowded areas and remove hanging branches. Respread rest of bottom tier of
branches with 1 m spreaders
Late May Chemically thin then follow up with hand-thinning to appropriate levels to ensure
regular annual cropping and adequate fruit size
July Spread second-tier branches with 30 cm nail-tipped spreaders where necessary
shape
Continued
350 T.L. Robinson
Sixth year
Dormant Head leader 1 m above established second tier to establish third-tier scaffolds 1 m
above second tier. Lightly dormant-prune to invigorate weak areas, thin out
overcrowded areas and remove hanging branches. Reposition bottom-tier spreaders
where necessary
shape
Seventh year
Dormant DO NOT HEAD LEADER. Remove all shoots competing with the leader, defining
permanent third-tier scaffolds. Remove largest one or two branches between first-
and second-tier scaffold limbs
shape
Mature-tree pruning
Dormant Limit tree height by cutting back into 2–3-year-old wood. Remove large branches
between tiers of scaffolds. Remove least desirable bottom-tier scaffold so that no
more than four limbs remain. Prune back lower-tier scaffolds to a side-branch to
facilitate movement of equipment and preserve fruit quality. Second- and third-tier
branches can be removed if excessively vigorous. During dormant-pruning of the
12th to 15th year, remove second-tier east- and west-growing branches to create a
palmette or flat fan-type top
August Summer-prune off vigorous upright shoots to increase light penetration and maintain
fruit colour. Occasionally tuck upright shoots where additional fruiting wood is
needed
As with the central-leader system, the 15.3.3 Palmette-leader system

canopy is developed by heading the leader
annually until the first two tiers are devel- The palmette-leader (PL) system is a modifi-
oped and by manually spreading the cation of the central-leader system that was
branches in each tier to an angle of 30° developed by Lakso et al. (1989a), which is
above horizontal for the bottom tier and to designed as a conversion form for mature
horizontal for the second tier. Above the central-leader trees. In many cases as central-
second tier of branches no permanent leader trees age, the upper limbs outgrow
branches are allowed to develop, but small the lower tier, resulting in excessive shade in
fruiting branches on the leader are retained the bottom of the trees. There is a strong ten-
and allowed to crop and bend below hori- dency for the upper scaffold limbs to grow
zontal. The bottom tier of branches is more vigorously than the lower limbs, due
developed at 60–90 cm above the ground to better light exposure. The shading that
but has only four branches. The second tier develops reduces flowering and fruiting in
of branches is about 1 m above the first tier the centre of the tree.
and also has four branches. After the sec- The PL shape was designed to improve
ond tier of branches is developed, the the light distribution in the tree canopy. This
leader is left unheaded and allowed to tree is developed by removing upper east-
fruit. After the leader reaches the desired and west-growing branches of a mature cen-
mature tree height, the leader is cut annu- tral-leader or mini-central-leader tree, result-
ally to a side-branch at about 3–3.5 m. ing in a flat north–south-orientated palmette
Second-tier branches are shortened when (i.e. fan-shaped) top. The lower tier or tiers of
they become too long to maintain the conic limbs are left intact. The primary advantage
shape. of the PL form is that large gaps are created
in the upper east and west sides of the The tree is a narrow, fully dwarf, conic-
canopy, which guarantee good light exposure shaped tree (Fig. 15.2), which allows planting
to all parts of the tree. A more subtle benefit of very high tree densities, ranging from 1500
is that good light penetration into the tree to 4000 trees ha1 in either single-, double-,
centre is found throughout the season. A use- triple- or multiple-row beds. The most com-
ful feature of the palmette top leader is that it mon rootstock used is M.9, but other stocks
can be adapted to east–west-orientated rows. of the same dwarfing level, such as B.9 and
With east–west-planted rows, the flat fan- G.16, are also used. The trees are always sup-
shaped top of the PL trees may still be orien- ported with either an individual tree stake or
tated north–south (i.e. perpendicular to the a single- to three-wire trellis. The width of the
row direction). This creates a gap between canopy is typically less than 2 m and tree
trees in the row for light penetration to the height ranges from 2 to 3 m. In Germany,
lower limbs of the canopy. mature tree height of slender-spindle trees
This tree form is most useful as a conver- was related to row spacing by dividing row
sion form for semi-dwarf to semi-vigorous spacing by 2 and adding 1 m (Winter, 1981).
central-leader trees between the ages of 7 The slender-spindle tree typically has a
and 15 years old. The conversion process is single permanent tier of branches, which are
done over a 3-year period by removing two developed by heading an unbranched tree
or three upper limbs each year. When trees (whip) at planting or by planting a branched
are between 7 and 15 years of age, the upper- tree (feathered) from the nursery (Table 15.2).
tier limbs are not large enough for the After planting, the central leader is cut at
removal of two or three upper-tier east- or 30 cm above the highest side-shoot. The
west-growing branches per year to result in feathers or lateral branches, which develop
excessive pruning and reduced yield. With during the first year, form a permanent tier
more severe pruning, yield is reduced exces- of branches. They are tied horizontal during
sively. The PL system results in improved the first or second year to induce cropping
light distribution to the bottom of the tree
and thus improved fruit quality. With older
trees, very large limbs must be removed to 2.25 m
make the conversion to PL and care should
be taken to make the conversion slowly over
several years so that excessive vigour and a
reduction in yield are not induced. Pruning
of this tree form is simplified compared with
that of the central-leader tree. Pruning con-
sists of keeping the palmette top narrow,
thinning out excess limbs in the top and
removing upright growth from the bottom
tier of scaffolds.
15.3.4 Slender-spindle system
The slender-spindle tree was developed by

Wertheim (1968) in the early 1960s and was
designed to improve early yields and man- 1.5 m
agement efficiency by planting higher tree Fig. 15.2. The slender-spindle tree has a supported
densities and reducing tree height to allow zigzagged trunk to limit tree height, a conic shape
all management to be done from the ground. and a permanent basal tier of branches. Lower
This system has become the dominant plant- branches are shortened to keep the tree slender
ing system in many parts of Europe while upper branches are shortened or removed and
(Oberhofer, 1987; Wertheim et al., 2001). replaced to maintain the conic shape.
352 T.L. Robinson
Table 15.2. Simplified pruning and training plan for the slender-spindle system. (NB. Northern-
hemisphere dates.) (Adapted from Robinson and Hoying, 1994.)
First year
At planting Plant highly feathered trees. Adjust graft union to 10 cm above soil level. Remove all
scaffolds below 60 cm, using a flush cut. Trees with three or more scaffolds (25 cm
long) should be headed at uppermost scaffold. The uppermost side-branch should
then be tied up as the new leader. Scaffolds longer than 45 cm should be headed by
removing one-third their length
5–10 cm growth Attach clothes-pegs to new side-shoots on leader to promote wide crotch angles
Early summer Install tree-support system that will allow tree to be supported to 2.5 m. Attach tree to
Second year
Dormant Where growth has been good (> 45 cm of terminal growth), tie leader and vigorous
scaffold branches horizontal. Alternatively, remove leader down to a horizontal
scaffold branch and tie it up as the new leader
Mid-June Tie leader back to stake, allowing smoothly curving bends to remain. Hand-thin fruits
to 15 cm apart
August Tie up lower scaffolds not expected to support the crop
Third year
Dormant Where previous leader bending has significantly weakened the top of the tree, DO
NOT PRUNE. Where leader is vigorous, tie leader horizontal or alternatively remove
leader down to a suitable horizontal side-branch and tie it up as the new leader
Mid-June Tie leader back to stake, allowing smoothly curving bends to remain. Tie down
vigorous limbs that will not bend with the weight of the crop. Hand-thin fruits to
15 cm apart
August Tie up lower scaffolds not expected to support the crop. Alternatively, do not tie up
but prune back scaffolds to prevent limb breakage and preserve tree structure
Fourth year
Dormant DO NOT PRUNE LEADER. Remove overly vigorous upright limbs that are more
than two-thirds the diameter of the leader, using a bevel cut. Tie down other
vigorous, upright limbs below the horizontal overlooked during third summer
Mid-June Tie leader to stake, allowing smoothly curving bends to remain. Tie up fruitful
scaffolds that will not support crop weight
shape. Tie up lower scaffolds not expected to support the crop
Fifth and sixth year

Dormant Limit tree height by cutting leader into 2- or 3-year-old wood back to a horizontal
fruiting branch. In each year, remove at least one undesirable bottom-tier scaffold
until four remain. Shorten bottom-tier scaffolds by pruning back to side-branch to
facilitate equipment movement and preserve fruit quality on lower limbs. Shorten
branches that have become pendant back to horizontal portion of the branch
shape
Mature-tree pruning
Dormant Limit height by cutting leader back to a fruitful side-branch. Shorten bottom-tier
scaffolds by pruning back to side-branch to facilitate equipment movement and
preserve fruit quality on lower limbs. Shorten branches that have become pendant
back to horizontal portion of the branch. Remove one or two vigorous upper-scaffold
limbs each year, preserving all weak-fruiting wood and permanent lower-tier scaffolds
August Summer-prune to encourage light penetration and maintain pyramidal tree shape
and to limit the width of the canopy. The develops and shading results (Robinson et al.,
leader is trained in a zigzagged manner up 1989). This is especially a problem with vig-
the support pole, either by annually remov- orous growth, where these gaps can be closed
ing the leader down to the first side-branch very quickly in the season, leading to poor
and tying up the lateral branch as the new fruit colour and quality if the trees are not
leader or by bending the leader to 45° from summer-pruned (Sansavini et al., 1981;
the vertical each year and then annually Corelli and Sansavini, 1989). The slender-
tying it back and forth across the pole (Plate spindle system has been most successful in
15.3). These procedures limit the growth of more northern climates where natural tree
the leader, which allows tree height to be vigour is less. In more southern climates, nat-
limited to 2.5–3 m. The upper part of the tree ural tree vigour on M.9 is often excessive,
is composed of small fruitful branches, which makes it difficult to manage trees at the
which bend with crop below the horizontal. very high tree densities of slender-spindle.
In the 1970s, most slender-spindle orchards
had densities from 1500 to 2500 trees ha1 and
had a tree height and diameter (i.e. spread) of 15.3.5 North Holland spindle
about 2 m. During the 1980s, higher tree densi-
ties of between 2500 and 3500 trees ha1 were The North Holland slender-spindle system
common and a narrower and taller tree form was developed by a Dutch fruit grower, D.
was developed, with a tree diameter of Huisman, in the Northern province of the
1.25–1.5 m and a tree height of 2.5 m. In the country in the late 1970s (Wertheim, 1981). It
1980s greater emphasis was placed on obtain- is a modification of the slender-spindle
ing significant yield in the second year after system that is more slender (1–1.5 m diameter)
planting by using feathered trees, where the and utilizes narrower in-row and between-
feathers started at 50 cm above the soil. The row spacings. Typically the North Holland
low height of the feathers required significant spindle is planted in a three-row bed. This
labour to tie them up when they began to fruit results in higher tree densities than the tradi-
to prevent fruit from touching the ground. In tional slender-spindle orchards. The trees are
the late 1990s, the minimum height of feathers planted on M.9 and are limited to a 2 m
was raised to 80–90 cm (Balkhoven-Baart et al., height. The tree is developed in much the
2000). This eliminated the need for tying up same way as a slender-spindle tree, but the
branches. In addition, many growers have lateral fruiting branches are cut back rather
allowed tree height to increase to 2.75–3 m drastically to keep the tree slender and small.
(Mantinger, 2000). In the 1990s, many slender- The North Holland spindle also differs from
spindle growers began to avoid pruning trees the slender-spindle in having a reduced
after planting or during the first few years. If amount of renewal pruning of lateral fruiting
the central leader was cut to the highest side- branches. This tree at maturity has a lower tier
shoot each year, a vigorous frame developed, of three short, horizontal, permanent scaffold
which needed a lot of summer-pruning labour branches at a height of 70–80 cm on the
to maintain good light distribution in the tree leader. Above the lower tier there are numer-
for good fruit quality. Without pruning of the ous shortened, semi-permanent fruiting
leader and with feathers starting at 80 cm, the branches along a zigzagged leader. The
tree can be allowed to crop in the second year, greater severity of pruning used with the
which gives natural bending of lateral North Holland spindle compared with the
branches to keep the canopy narrow slender spindle was intended to improve fruit
(Mantinger, 2000). quality by improving light exposure to the
The narrow, slender shape of the slender- lower and centre portions of the canopy. The
spindle canopy helps ensure that most of the system has worked very well in low-vigour
canopy is well exposed. However, in some soils or climates. With more vigorous soils or
cases, if the branches are too close together climates, the severe pruning has stimulated
within the canopy with only small gaps too much growth, which has resulted in
between them, a high density of foliage poorer fruit quality than other systems.
354 T.L. Robinson
15.3.6 Slender-spindle multi-row or bed M.26. Trees are supported with a single- to
system multiple-wire trellis up to 3 m high. The verti-
cal-axis tree is made up of a single vertical
The slender-spindle multi-row or bed system trunk, which serves to support many rela-
was developed in Holland in the 1970s. It tively small-diameter fruiting branches (Fig.
is a modification of the single-row slender- 15.3 and Plate 15.4). Maintaining the integrity
spindle system that eliminates the tractor of the apical bud of the leader during tree
alleys, leaving only a walking path between development is important for ensuring the
the rows. This results in very high tree densi- development of weak fruiting branches
ties. The beds can be from three to seven (Lespinasse and Delort, 1986). At planting
rows wide, with a tractor alley between beds and during the development years, the leader
(Wertheim, 1978b, 1981; Oberhofer, 1987). In is not headed back (Table 15.3). The weak-
a few experimental cases, tractor alleyways growing fruiting branches develop along the
were eliminated completely and a full field trunk in a manner dependent on the fruiting
system was planted and managed with over- type of the cultivar. Fruiting branches are
the-row equipment (Wertheim, 1981). The retained along the trunk from a height of 1 m
trees are planted on M.9, or in some cases, upward. For trees on M.9, a good balance
the super dwarfing M.27 rootstock. Spacing between vegetative growth and fruiting is
between trees and between rows is similar usually obtained with 12 to 16 fruiting
(1.0–1.75 m). Mature tree height is 2 m and branches (Lauri and Lespinasse, 2000). The
the tree canopy resembles that of a slender- number and distribution of branches are con-
spindle tree. The trees are supported by indi- trolled by pruning. As the tree matures, fruit-
vidual tree stakes. With most slender-spindle ing branches are periodically renewed after
bed systems, the between-row spacing in the bending under the weight of the fruit, and
beds is less than the in-row spacing and trees hence do not become permanent scaffold
are planted on a diagonal pattern, so that branches (Plate 15.5). Heavy fruiting controls
mini-rows are created diagonally across the
beds. The diagonal alleyways across the beds
4m
aid in spray penetration to the interior of the
beds, as well as providing picker access. The
multi-row bed systems achieved very high
production and light interception by the sec-
ond or third season in the orchard (Palmer et
al., 1992). As the trees matured, shading
between trees often resulted in reduced fruit
quality and increased summer-pruning costs
(Balkhoven-Baart et al., 2000).
15.3.7 Vertical-axis system
The vertical-axis system was developed in the

late 1970s by Lespinasse (1980) in southern
France. Lespinasse (1977) categorized apple
cultivars into four types, based on fruiting
and growth habit. The vertical-axis system
was designed to utilize the natural fruiting 22 m
m
and growth habit of each cultivar. The verti- Fig. 15.3. The vertical-axis tree has a supported
cal-axis system utilizes tree densities from dominant trunk, a conic shape, a permanent basal
1000 to 2500 trees ha1, spaced at 1.0–2.0 m in tier of branches and small-caliper upper fruiting
the row and 4.0–5.0 m between the rows. The branches. Upper branches are shortened and then
most commonly used stocks are M.9 and removed and replaced to maintain the conic shape.
Table 15.3. Simplified pruning and training plan for the vertical-axis system. (NB. Northern-hemisphere
First year
at 110 cm above the soil line, with all scaffolds headed by removing one-third their
length. Trees with fewer than three feathers should be headed at 90 cm and all
feathers removed, using a bevel cut, leaving a 2 cm stub
Second year
Dormant DO NOT HEAD LEADER OR PRUNE TREES. If additional scaffolds are needed,
score above appropriate trunk buds at bud break
10–15 cm growth Pinch lateral shoots in top one-quarter of last year’s leader growth, removing about
5 cm of growth (the terminal bud and four or five young leaves)
Mid-June Repinch all lateral shoots in top one-quarter of last year’s growth. Tie developing
leader to support system with permanent tie. Remove all fruit on 1-year-old wood
and hand-thin remaining fruits to 15 cm apart
Mid-July Repinch vigorous lateral shoots in top one-quarter of last year’s growth. Tie down
four or five permanent lower scaffold branches to the horizontal. Attach permanent
trellis clips to tree to support fully second-year crop
Third year
Dormant DO NOT PRUNE LEADER. Remove overly vigorous upright limbs that are more
than two-thirds the diameter of the leader, using a bevel cut. Tie down other vigor-
ous, upright limbs below the horizontal overlooked during second summer
10–15 cm growth Pinch lateral shoots in top one-quarter of last year’s leader growth removing about
5 cm of growth (the terminal bud and four or five young leaves)
June 15 Repinch all lateral shoots in top one-quarter of last year’s growth. Tie developing
leader to support system with a permanent tie. Hand-thin to single fruits spaced
10 cm apart
Mid-July Repinch vigorous lateral shoots in top one-quarter of last year’s growth
but prune back scaffolds to prevent limb breakage and preserve tree structure
Fourth year
Dormant DO NOT HEAD THE LEADER. Remove overly vigorous upright limbs that are more
ous, upright limbs below the horizontal overlooked during third summer
Late May Chemically thin, then follow up with hand-thinning to appropriate levels to ensure
shape. Tie leader to support system with a permanent tie at the top of the pole

Dormant DO NOT HEAD THE LEADER. Shorten bottom-tier scaffolds where needed back to
side-branch to facilitate movement of equipment and preserve fruit quality on lower
limbs. In each year remove one of the least desirable lower-tier scaffold branches
until only four remain. Shorten branches that have become pendant back to horizon-
tal portion of the branch. Remove up to one vigorous upper-scaffold limb each year
to begin renewal of fruiting branches
Continued
356 T.L. Robinson
Fifth and sixth year—Continued

Late May Chemically thin, then follow up with hand-thinning to appropriate levels to ensure
shape
Mature-tree pruning
Dormant Limit tree to desired height by cutting leader back to a fruitful side-branch. Shorten
bottom-tier scaffolds by pruning back to side-branch to facilitate equipment move-
ment and preserve fruit quality on lower limbs. Remove one or two vigorous upper-
scaffold limbs each year, preserving all weak-fruiting wood and permanent lower-tier
scaffolds
vegetative growth in the top, guaranteeing a American growers in the 1990s and has now
uniform and moderate growth vigour in all become the dominant orchard system in
parts of the tree. This helps maintain a nat- many parts of the world; however, many
ural equilibrium between fruiting and vegeta- growers have made modifications to fit their
tive growth. The growth of the leader is commercial orchards (Granger and Philion,
naturally weakened by fruiting in the third 1988; Maillard and Herman, 1988; Rice, 1988;
and fourth years, causing it to bend under the Tukey, 1988; Barritt, 1992; Perry, 1994, 1998;
weight of fruit, limiting its height. By the time Barden, 1995; Quamme and Geldart, 1995;
this happens, trees have usually reached a Robinson et al., 1996a; Karasiewicz, 1997).
height of 3–3.5 m. The primary modification from the original
Lespinasse and Delort (1986) identified vertical-axis protocol of Lespinasse has been
three fruiting zones in fruiting limbs. ‘Zone the development of a permanent tier of
A’ (0–30° from the vertical) is characterized lower scaffold branches, which are not
by the development of strong vegetative renewed. Other modifications include pinch-
shoots, which are able to bear fruits of good ing of vigorous laterals to suppress their
quality 1–2 years later. ‘Zone B’ (30° and 120° growth, shortening of fruiting branches to
from the vertical) is the prime fruiting zone, reduce sunburn, bending of the leader to
with moderate shoot growth. ‘Zone C’, reduce leader vigour, and replacing leaders
which is below 120°, produces smaller infe- with reproductive laterals on vigorous-grow-
rior fruits because of the deleterious effects ing trees (Perry, 2000). While these modifica-
of within-tree shading on fruit quality. The tions have been introduced, the core of the
vertical-axis training system aims to develop original vertical-axis protocol is still followed
limbs in zone B, which produce the best fruit by many growers today.
quality. When a fruiting branch develops, it
usually has a vertical orientation (zone A). If
it is left unpruned, generally it will bend into 15.3.8 SolAxe system
zone B, with heavy cropping in the second or
third year. If the branch becomes too pen- In the mid-1990s, Lespinasse (1996) devel-
dant (zone C) from repeated fruiting, it is oped a modification of the vertical-axis,
removed and a renewal branch is developed called the SolAxe, by focusing even more
in its place. This system of pruning, there- on branch bending and avoiding renewal
fore, keeps fruiting on relatively young pruning in an attempt to gain a more weep-
organs, i.e. 2- to 3-year-old spurs and ing canopy. Studies by Lauri and
crowned brindles. Lespinasse (1993) showed that the poor
The vertical-axis system was embraced by fruit characteristics of branches in zone C
French growers in the 1980s and by North are due to the shading caused by other
branches kept above them. Additionally the 15.3.9 Slender-pyramid system

A, B, C zone concept does not fit the termi-
nally bearing cultivars (type IV, according The slender-pyramid system was developed
to Lespinasse, 1977), such as ‘Granny by Tustin and colleagues in the late 1980s in
Smith’ or ‘Rome Beauty’. In these cultivars, New Zealand (Tustin et al., 1990; Tustin, 2000).
the growing shoot generally ends in a fruit- In designing their system they considered
bud and is able to develop a bourse shoot, physiological principles of light interception,
which will fruit again the following year. light distribution and pruning, plus combined
To utilize this natural balance between veg- features of the vertical-axis, McKenzie central-
etative growth and fruiting on type IV cul- leader and slender-spindle systems
tivars, they suggested a new (McKenzie and Mouat, 1963; Wertheim,
fruiting-branch training concept, which did 1978a,b; Lespinasse and Delort, 1986). The
not utilize removal of pendant shoots back slender-pyramid management system can be
to more vertical renewal branches, but used with relatively low tree densities (600
relied on the removal of the vertical trees ha1) with MM.106 rootstock and with
renewal shoots to keep the pendant fruiting moderately high tree densities (2000 trees
branch well exposed and productive. The ha1) with M.9. The slender-pyramid tree has
name of the new system, SolAxe, was taken a dominant central leader, a well-developed,
from the ‘Solen’ system, on which the new spreading, basal tier of four branches bearing
fruiting-branch concept was initially devel- fruiting laterals, with a narrow pyramidal
oped during the 1980s, and from the verti- canopy above the basal tier (Fig. 15.5). The
cal-axis system. The system combines the mid- and upper canopy is comprised of well-
bending of the central axis and the fruiting spaced whorls of weaker fruiting laterals
branches from the Solen system with the arising directly off the trunk. Trees are not
free-growing fruiting branches and the headed when planted and are supported
removal of competing vegetative branches using a single- or multiple-wire vertical trel-
from the vertical-axis system. lis (Table 15.4). Unrestricted extension
The tree is developed at planting by growth of the central leader is encouraged by
strongly bending the feathers along the removal of competing lateral shoots early in
trunk into an arc. Branches that are below the summer each year.
1.2 m are removed. The leader is attached to
a support system either in the autumn of
the first growing season or at the end of 3.5 m
spring in the second season. The leader is
left unheaded and allowed to grow to the
top of the support system (Plate 15.6).
When the leader reaches the top of the sup-
port system, it is bent horizontal, either nat-
urally by fruiting or artificially by bending
(Fig. 15.4). Upper fruiting branches are
bent below the horizontal, either naturally
through the weight of the fruits or artifi-
cially by tying during the second or third
year. Excess branches are removed by prun-
ing, beginning in the fourth year. As with
the vertical-axis system, the best balance
between vegetative growth and fruiting on 2m
a mature SolAxe tree is obtained with 12–16 Fig. 15.4. The SolAxe tree has a supported
fruiting branches arranged spirally along dominant trunk, with all branches trained to a
the trunk. On each fruiting branch, upright pendant position. Branches are not renewed. The
shoots are removed annually and the pen- upper part of the tree is bent horizontal to limit tree
dant portion is retained. height and reduce vegetative growth in the top.
358 T.L. Robinson
4m mum of four, arranged in a cruciform array

across and along the row. Mature tree man-
agement utilizes renewal pruning for
replacement of fruiting branches that are too
vigorous or long to maintain a satisfactory
light environment within the canopy.
15.3.10 HYTEC system
The hybrid tree cone orchard system

(acronym HYTEC) was developed in the late
1980s by Barritt (1991, 1992) specifically for
arid apple-producing areas that have signifi-
cant fruit sunburn. The system also has the
goals of high, early and sustained yield per
hectare, high labour efficiency and high fruit
quality. It is a blend of the slender-spindle
2.5 m
(Wertheim, 1978b) and vertical-axis
Fig. 15.5. The slender-pyramid tree has a supported
(Lespinasse, 1980) systems that combines
dominant trunk, a conic shape and well-defined
nuances of both systems and is intermediate
tiers of branches. A hierarchy of branch thickness is
maintained from the bottom to the top of the tree by in canopy height between the two (Barritt,
removing large-diameter branches in the upper part 1992). In addition, techniques for vigour con-
of the tree. Conic shape is maintained by shortening trol and minimal pruning used with angled-
upper branches. canopy systems, such as the Tatura trellis,
the Geneva Y trellis and the Güttinger V, are
also incorporated into central-leader man-
In the first year, the growth of six to eight agement of HYTEC trees. To reduce vigorous
basal-tier limbs is encouraged by early- growth high in the tree (often a feature of
summer removal of unwanted competing mature vertical-axis trees), the central-leader
lateral shoots from the central leader. No of a HYTEC tree, during the formative years,
further pruning in the first year is required. is pruned and/or bent annually in a manner
During the second year, basal tier limbs are similar to that of the slender-spindle system.
spread down to 15–20° above the horizontal. To achieve greater production at full canopy
Extension of the basal limbs is encouraged than slender-spindle trees, the HYTEC tree is
by ensuring that there is only a single apical taller with greater canopy volume, a charac-
shoot on each limb. In the second and third teristic of vertical-axis trees. Light distribu-
years, selection and thinning among lateral tion is maintained in the HYTEC tree by
shoots arising from the central leader above maintaining an open canopy structure. Fruit
the basal tier are done in early summer. In sunburn is also reduced on HYTEC trees by
the second year, fruiting is allowed only providing some limb-to-limb shading
when well-branched nursery trees are used through stiffening lateral limbs with shorten-
and then only on spurs on the basal-tier ing pruning, a technique used with slender-
limbs or directly on the central leader. This spindle training, but not in the training of
system results in rapid canopy development, vertical-axis trees (Fig. 15.6).
with a hierarchy of fruiting branches of flat Mature trees are 3 m tall and cone-shaped,
orientation and decreasing vigour up the with a central leader and a basal width of
central leader (Plate 15.7). From the third 1.5–2.25 m. Tree density is from 1400 to 2300
year, cropping increases rapidly, which limits trees ha1, with spacing from 1.25 to 2.0 m
continued expansion of the tree canopy. As between trees in the row and 3.5 to 4.25 m
the tree matures, the number of limbs in the between rows. Each tree is supported with
basal tier is reduced progressively to an opti- either a three-wire vertical trellis, an individ-
Table 15.4. Pruning and training plan for the slender-pyramid system. (NB. Northern-hemisphere dates.)
(Adapted from Tustin, 2000.)
First year
using a flush cut. DO NOT HEAD THE LEADER
5–10 cm growth Remove vigorous shoots that are competitive with the leader
Second year
Dormant DO NOT HEAD THE LEADER. If additional scaffolds are needed, score above
appropriate trunk buds at bud break
Early summer Limit cropping to spurs close to the base of the basal-tier limbs
Mid-July Tie down four to six permanent basal-tier scaffold branches to 15° above horizontal
Third–fifth year
than two-thirds the diameter of the leader, using a bevel cut
Early summer Remove lateral shoots that are excessively vigorous, have narrow crotch angles or
are in poor positions. The branches along the leader of the tree should have a hier-
archy of decreasing vigour up the leader
Fourth–sixth year
than two-thirds the diameter of the leader, using a bevel cut. Progressively reduce
the number of basal-tier branches to an optimum of four arranged in a cruciform
array across and along the row, by removing one each year
Early summer Remove lateral shoots that are excessively vigorous, have narrow crotch angles or
are in poor positions. The branches along the leader of the tree should have a hier-
archy of decreasing vigour up the leader
Mature-tree pruning
Dormant Remove and renew upper-scaffold limbs when they cause excessive shading and to
develop new fruiting wood. Limit tree to desired height by cutting leader back to a
fruitful side-branch. Shorten bottom-tier scaffolds by pruning back to side-branch to
facilitate equipment movement and preserve fruit quality on lower limbs
ual post or a thin bamboo or metal conduit in a manner similar to slender-spindle tree
pole secured to a single-wire-trellis support training. The lower tier of horizontal scaffold
system. The support is used to train the tree limbs is permanent. Scaffold branches are
vertically and to prevent tree leaning and/or shortened to fit within their allotted space by
breakage during strong winds and with heavy pruning to a weak lateral or spur. Upper limbs
crops. The most common rootstocks for non- are trained horizontally and, after bearing
spur cultivars are M.9, B.9 and M.26 (Barritt et fruit, are shortened into older wood to retain
al., 1995). Annual pruning or bending of the the tree’s cone shape, to stiffen limbs and to
central leader – a decision based on central- stimulate shoot growth, which provides tran-
leader vigour – encourages the development sient shade that reduces fruit sunburn.
of strong lower-tier scaffold branches, Central-leader height is limited to 3 m by
stimulates branching and reduces tree height pruning into older wood to a weak lateral.
360 T.L. Robinson
3m cally; or (iii) left unpruned and bent at a 45°

angle (Barritt, 2000) (Plate 15.8). Heading
can be used if central-leader growth is weak
(generally less than 50 cm in length), to
encourage stronger growth and stimulate
branching. At intermediate vigour levels
(between 50 and 100 cm of growth), the cen-
tral leader can be left unpruned as there is a
need neither to stimulate branching by
heading nor to reduce vigour. With exces-
sive vigour levels (generally more than
100 cm in length), bending of the central
leader is used to reduce tree height and to
stimulate branching. Bending or zigzagging
the central leader is a hybrid technique that
2m falls between the vertical-leader training of
the vertical-axis system and the angled
Fig. 15.6. The HYTEC tree has a supported
zigzagged trunk, a conic shape and a permanent
leader of systems such as the Tatura trellis,
basal tier of branches. Upper branches are the Geneva Y trellis or the Güttinger V.
shortened to stiffen them to prevent movement with Angling the leader at 45° reduces extension
fruit load. Large upper branches are removed. growth; however, the whole tree is still ori-
entated vertically. The zigzag technique
reduces tree-height extension and stimulates
At planting, unbranched trees (whips) are lateral branching. Until the tree reaches its
pruned at 80–90 cm above the soil and final height of 3 m, a decision about central-
branched (feathered) trees are pruned at leader training must be made each dormant
about 25 cm above the top useable branch, season. The decision is based on the vigour
which is usually 90–115 cm above the soil of the central leader. Therefore, it may be
(Table 15.5). Branches that are longer than necessary to bend the central leader in one
about 50 cm are headed lightly up to one- year if it has had very strong growth, but in
third their length to stiffen the branches, to the next season, if growth is moderate, the
reduce blind wood and to reduce top : root appropriate central-leader-training tech-
ratio and thus lessen transplant shock. nique will be to tie it vertically and avoid
Branches are trained to an angle between any other intervention. When the tree
horizontal and 30° above the horizontal. exceeds a height of 3 m, the leader is cut
The HYTEC system utilizes pruning back into older fruiting wood to a weak hor-
and/or bending techniques for training the izontal branch.
central leader to ensure adequate branching The lower-tier scaffold limbs are perma-
along the leader and to control vigour in nent and are trained horizontally or at a slight
the top of the tree. In some climates, an upward angle (< 30°). Other lateral branches
unpruned central leader produces excessive are trained horizontally and are maintained
growth and insufficient lateral branches. The within their allotted space by shortening into
few lateral branches that develop are below older wood, usually to a weak lateral or spur,
the terminal bud and are narrow-angled either in the dormant season or during the
shoots. A 0.5–1 m section of the central summer. Shortening pruning is used to stiffen
leader may be without lateral branches. limbs. Stiff limbs have less fruit sunburn than
With the HYTEC system, it is essential to flexible limbs, which change position during
assess central-leader vigour before deciding the season as fruit weight increases. The addi-
on the appropriate management procedure. tional branching induced behind the shorten-
The central leader of HYTEC trees can be ing cut also provides temporary (transient)
either: (i) lightly or moderately pruned by shade, which reduces fruit sunburn. Limbs
heading; (ii) left unpruned and trained verti- are usually shortened following their third
Table 15.5. Pruning and training plan for the HYTEC system. (NB. Northern-hemisphere dates.)
(Adapted from Barritt, 2000.)
First year
at 110 cm above the soil line. Branches longer than 50 cm should be headed by
removing one-third their length. Trees with fewer than three feathers should be
headed at 90 cm and all feathers removed, using a bevel cut, leaving a 2 cm stub
Second year
Dormant Bend leader to a 45° angle and attach to the support pole
Mid-July Tie down four or five permanent lower-scaffold branches to slightly above the
horizontal
Third year
Dormant Bend leader to a 45° angle in opposite direction of the bend from the previous year
and attach to the support pole. Remove overly vigorous upright limbs that are more
ous, upright limbs below the horizontal overlooked during second summer
August Shorten lower-scaffold branches to allotted tree spacing and to prevent limb
breakage to preserve tree structure
Fourth–sixth year
Dormant Bend leader to a 45° angle in opposite direction of the bend from the previous year
and attach to the support pole. Remove overly vigorous upright limbs that are more
ous, upright limbs below the horizontal overlooked during third summer. Shorten
horizontal fruiting branches to stiffen them and limit their length
shape
Mature-tree pruning
Dormant Limit tree to desired height by cutting leader back to a fruitful side-branch. Shorten
bottom-tier scaffolds by pruning back to side-branch to facilitate equipment move-
ment and preserve fruit quality on lower limbs. Shorten upper branches to maintain
cone shape. Periodically remove an upper branch that has become too large and
vigorous
year of growth and may be shortened again in 2000). If shading of the lower canopy reduces
subsequent years. Eventually upper limbs are flowering, fruit set, fruit size and colour to
completely removed. The decision to remove unacceptable levels, limbs higher in the tree
a limb is based on the amount of shading it must be removed, regardless of their diame-
causes lower in the tree canopy and its vigour. ter. A short stub is retained as a site for a new
To improve canopy openness and light distri- replacement shoot. At maturity, a row of
bution, limbs are usually removed when their HYTEC trees, when viewed from the side,
diameter is 50% of the diameter of the central should have a sawtooth pattern between tops
leader at the point of attachment (Barritt, of adjacent trees.
362 T.L. Robinson
Once a HYTEC tree reaches full canopy tree diameter. The slender-spindle has a
size, maintenance pruning includes: (i) short- diameter of 1.0–2.0 m, the super spindle 0.5
ening pruning of the central leader (to limit to 1.0 m and the string tree less than 0.5 m.
tree height), the upper branches (to maintain Most commercial super-spindle orchards
the cone shape) and the permanent lower have a row spacing of 3 m or less and a
scaffold branches (to limit tree width); (ii) planting distance within the rows of
periodic removal of upper limbs to renew 45–80 cm, giving a density of 4000 to 7500
fruiting branches and improve sunlight dis- trees ha1. Row distances less than 3 m can
tribution; and (iii) removal of unwanted be too narrow to obtain high picking out-
shoots, including forked branches and vigor- puts. Mature tree height is 2.5–2.75 m.
ous upright shoots. The super-spindle system has a goal of
some production during the first year in the
orchard. This can best be achieved with
15.3.11 Super-spindle system nursery trees that have some flower buds
(generative trees). Often growth-regulator
The modern version of the super-spindle treatments of multiple low-dose sprays of
system or cordon system was developed in naphthaleneacetic acid (NAA) or ethephon
southern Germany by Nüberlin in the early are applied in the nursery row to stimulate
1990s (Weber, 2000). However, the fruit-wall flower-bud production of lateral buds on
system developed by Luckwill (1978) at 1-year wood. Super-spindle orchards do
Long Ashton research station was an earlier not usually use the highly branched and
version of the super-spindle system that relatively expensive trees used in slender-
relied on the use of growth-regulating and spindle orchards, but rather whips with a
flower-regulating chemicals to limit tree size number of short shoots along the leader or
and induce heavy cropping. The German even cheaper trees, such as budded root-
super-spindle system is a modification of the stocks (sleeping-eye trees) (Weber, 1997).
slender-spindle system that utilizes very
high tree densities and draws upon the
concepts of the ‘Bleiber–Weicher’ planting 2.5 m
system, which was originally developed by
Dickenmann, a nurseryman in Switzerland.
That system suggested planting compact,
slim apple trees with spurs along the central
leader at very close spacings, in single rows
and then pulling out every second tree after
4 or 5 years once canopies became too
crowded. Nüberlin suggested leaving the
trees at the original planting distance
throughout the life of the orchard and devel-
oping management strategies to keep the
trees compact without the development of
thick lateral branches (Nüberlin, 1993; Fig.
15.7 and Plate 15.9).
The goals of the super-spindle system are
very early and high yields so that new culti-
vars can be introduced as quickly as possible
to meet market demands, less manual work,
less chemical input and high picking output, 0.6 m
resulting in low fruit cost per hour of labour. Fig. 15.7. The super-spindle tree has a supported
No exact definition exists of what a super- trunk, a narrow cylindrical shape and no permanent
spindle orchard is. Österreicher (1993) classi- branches. Vigorous branches are removed, leaving
fied the different systems according to their only small-diameter fruiting branches on the trunk.
Because of the very high total tree costs per 15.3.12 Meadow-orchard system
hectare due to the high tree densities of this
system, most growers grow their own trees The meadow-orchard system was developed
to reduce the cost of plant material. by Hudson (1971) and Luckwill (1978) as the
Super-spindle trees are trained differently ultimate in apple-orchard intensification with
from slender-spindle trees (Table 15.6). To trees planted at a density of 70,000 trees ha1.
reduce tree growth and maximize early fruit- The trees are spaced 30 cm 45 cm and have
ing, super-spindle trees are not headed at a tree height of 1 m. The orchard is managed
planting and are often planted on top of the like an agronomic crop, with all of the
ground and a small amount of soil is pushed machinery work done over the row. The sys-
up to cover the roots. In the first year, the tem is based on an alternate-year cropping
steep-angled and strong lateral branches schedule and mechanization of harvest. The
(larger in diameter than half the size of the orchard is developed by planting a budded
central leader) are removed by ripping them rootstock and, during the first year in the
from the tree. Growth-regulator sprays of orchard, treating the developing shoot with
NAA or ethephon can also be used to growth- and flower-regulating chemicals to
stimulate flower-bud initiation. In the second induce flowering. The tree is allowed to crop
year, no significant pruning is done. In the in the second year and produces from seven
third year, weak shoots are cut back into to 13 fruits. At harvest the entire top of the
2-year-old wood to promote shoot strength. tree including the fruit is cut off with a
Water sprouts are removed through ripping. mechanical harvester/combine and the fruit
In the fourth year, weak shoots are cut back is separated from the tree top. The 2-year
into 2-year-old wood and tree height is cycle is then repeated. Several rootstocks can
limited by one single cut into 2-year-old be used, such as M.26 and MM.106. Because
generative wood. of the very high tree density, this system is
The orchard life of super-spindle based on having very cheap trees, which
orchards can be as short as 7 years, but gen- would probably have to be propagated by
erally not longer than 12–15 years. This hardwood cutting and would be own-rooted.
means that the economic success of super- Very high experimental yields of 100 t ha1
spindle orchards depends to a large extent were achieved, but only every other year.
on very early, high yields of a high-priced This system was never planted commercially
new cultivar, low-priced trees from the nurs- with apples, but commercial versions were
ery, higher picking output and fewer man- established with peaches.
agement hours to maintain the system.
Fixed costs for the establishment of a super-
spindle orchard are higher than for other 15.3.13 Advantages and disadvantages of
conic-shaped systems
systems and must be justified by the market
returns of the cultivar and the early yields.
Conic-shaped systems are currently the
The high cost of the system makes it a
dominant tree form in commercial orchards
riskier system than more moderate-density
in most parts of the world. The primary
systems. However, if there is an economi-
advantages of the conic shape are as follows:
cally friendly market situation with a new
high-priced cultivar that has good fruit size 1. It is a natural tree form for apple that does
and a non-biennial bearing habit, coupled not require extensive branch and leader
with inexpensive plant material, profitabil- manipulation to produce and is easy to man-
ity for a new super-spindle orchard can be age. This allows trees to be developed with
achieved in a short time period. This per- minimal labour for tree training.
mits a short orchard lifetime, which gives 2. Its shape gives good light distribution
growers flexibility to respond to new culti- throughout the canopy by limiting the width
vars and changes in market demand of of the top of the tree. This results in minimal
existing cultivars. shading by the upper limbs of the lower limbs.
364 T.L. Robinson
Table 15.6. Pruning and training plan for the super-spindle system. (NB. Northern-hemisphere dates.)
(Adapted from Robinson and Hoying, 1994.)
First year
At planting Plant tree with as much of the rootstock shank out of the ground as possible, while
still covering all main brace roots, and tamp soil around roots. Remove all scaffolds
below 50 cm, using a flush cut. DO NOT HEAD LEADER. Prune off only diseased
and broken scaffolds. Provide trellis support immediately and tie tree to bottom wire
Late June Spray trees with low doses of NAA three times spaced at 2-week intervals, to
stimulate flower-bud formation for the second leaf
Second year
Dormant DO NOT HEAD LEADER. DO NOT PRUNE
Early June Hand-thin crop to single fruits 10 cm apart
Late June Spray NAA at 5 p.p.m. three times, spaced at 2-week intervals, to stimulate flower-
bud development for the third leaf. Attach developing leader to second wire
August Attach permanent trellis clips to tree to support fully second-year crop
Third year
Dormant DO NOT PRUNE LEADER
Late May Chemically thin according to crop load, tree strength and weather conditions, then
follow up with hand-thinning to appropriate levels to ensure regular annual cropping
and adequate fruit size
Late June Break vigorous shoots but leave attached to tree
August Prune out excessively vigorous shoots with a bevel cut. Lightly summer-prune to
encourage good light penetration and fruit colour
Mature-tree pruning
Dormant Rip out by hand excessively large and vigorous shoots (if diameter is > 50% that of
the leader), particularly in the top, where regrowth is not wanted. Prune out
excessively vigorous shoots (if diameter is > 50% that of the leader) in the bottom
with a bevel cut where new shoots are wanted. Shorten pendant shoots to a fruit bud
to keep the tree within its space, if needed
Late May Chemically thin according to crop load, tree strength and weather conditions, then
follow up with hand-thinning to appropriate levels to ensure regular annual cropping
and adequate fruit size
August Summer-prune to encourage good light penetration and fruit colour by cutting out
vigorous shoots and shortening back horizontal shoots to first fruit
The primary disadvantages of conic- wider than optimum row spacings to allow
shaped systems areas follows: the use of existing equipment. This has
resulted in relatively low yields for many
1. A substantial amount of light energy falls
dwarf orchards. Alternatively, if growers
between the tree rows on the tractor alley-
choose to plant wider than optimum row
ways and is wasted. This energy could be
spacing, they should grow taller trees, which
used to produce fruit. The only way to
capture more light.
reduce the wasted sunlight energy is to plant
2. As conic-shaped trees mature, the upper
rows very close together or to increase tree
branches of the tree begin to outgrow the
height. Close row spacings, such as are used lower branches, causing excessive shade on
in the meadow orchard or the super spindle, the lower canopy. Often the shape of mature
require growers to purchase new narrow trees resembled an inverted pear and eventu-
equipment. Many growers who want to ally trees took on an umbrella shape. The
plant dwarf trees are sometimes hesitant to central-leader system during the 1970s and
purchase new equipment and consequently 1980s had this problem when the upper tiers
choose to plant conic-shaped trees with of branches were considered permanent.
Systems that have emphasized renewal of of these were the espaliers grown along
upper branches, such as the vertical-axis and walls and fences. The first commercial apple
slender spindle, have been more successful orchards with restricted two-dimensional-
in maintaining the conical shape throughout plane canopies were developed by Italian
the life of the tree. fruit growers in the mid-1950s, which they
called the palmette training system. Their
Over the last 50 years the majority of
motive was to improve orchard labour effi-
apple growers in the world have opted to
ciency by the use of platforms, which
plant one of the conic-shaped systems. In
allowed more efficient picking, pruning and
the 1960s and 1970s, the central-leader sys-
thinning of the palmette trees compared with
tem was the most common system in the
the traditional vase-shaped fruit trees of that
world. However, in Europe, the slender
time. With the use of platforms to position
spindle and the vertical axis were the most
workers near their work (pruning and pick-
common systems in the 1970s and 1980s. At
ing), labour efficiency can be improved by
the end of the 20th century, the most com-
15–20%. The success of the palmette system
mon systems are the slender spindle and its
stimulated widespread grower adoption in
variants and the vertical axis and its vari-
Italy. Italian research with this system led to
ants. The slender spindle is most common
significant technical innovations in pruning
in northern Europe and the vertical axis or
to hasten fruit bearing and reduce pruning
one of its variants is most common in
costs over the life of the orchard and to
southern Europe, North America, South
increased planting densities to reap the bene-
America, New Zealand, Australia, South
fits of earlier and greater returns (Sansavini,
Africa and Japan. Over the last decade, the
1983). A number of variations of the palmette
height of most slender-spindle trees has system have been developed. Most of the
increased to close to 2.75 m and thus the newer palmette systems utilize dwarfing
trees are more similar to vertical-axis trees. rootstocks, which were not part of the origi-
This has allowed slender-spindle orchards nal palmette system. The suitability of the
to intercept more light and achieve higher palmette system to the use of picking plat-
yields, while reducing the amount of prun- forms helped it gain widespread acceptance
ing required to contain the tree to a short in Italy in the 1960s and 1970s, but, as the use
stature of 2 m. Likewise over the last of dwarfing apple rootstocks became com-
decade, the planting density of vertical- mon in the 1980s and 1990s, the advantage of
axis orchards has increased from 1500 to picking platforms disappeared.
more than 2000 trees ha 1, which is similar
to the density of slender-spindle orchards.
Thus the evolution of the two leading sys- 15.4.1 ‘Regular’ palmette system
tems in the world has resulted in great sim-
ilarities between the systems. In a few The original or ‘regular’ palmette training
locations in the world where land prices system was developed by Baldassari, a fruit
are very high or where there are subsidies grower in Ferrara, Italy, in the mid-1950s
from the government, the super-spindle (Sansavini, 1993). The tree is trained to a 3 m
system, with its very high tree densities, is tall four- to six-wire vertical trellis, which
being planted. serves to support the tree and crop at matu-
rity. The palmette tree has a central trunk
and regular tiers of branches that are trained
15.4 Flat Planar Canopy Systems to the two-dimensional vertical plane along
the row. Branches that are not in this plane
European gardeners have utilized trellises to are removed or trained over to the wire trel-
restrict apple tree canopies to a two-dimen- lis in the row plane. The branches at each tier
sional plane for centuries and have devel- are tied either at an oblique angle (30°), so
oped artistic apple trees of many geometric that they grow up across the wires (oblique
shapes (Huggard, 1980). The most common palmette) (Fig. 15.8), or horizontally along
366 T.L. Robinson
2.5 m style of pruning delayed the onset of produc-

tion because of the heading cuts. Later ver-
sions of the palmette system used feathered
trees (‘anticipated’ or ‘sprint’ palmette) from
the nursery, where the feathers could be
used to form the lower tier of branches with-
out the use of heading cuts.
15.4.2 ‘Free’ palmette system

2.5 m
The ‘free’ palmette system is a variation of
Fig. 15.8. The oblique-palmette tree has a central
the regular palmette system, but without its
trunk with four or five tiers of permanent scaffold
branches, which are trained at a 45° angle above
rigid structure, allowing the formation of
horizontal oriented along the row, forming an branches from laterals emerging from the
overlapping lattice structure between trees along the trunk without any geometric pattern. With
row. this system, heading cuts at planting and
thereafter are avoided, unless the tree needs
the wire at each tier (horizontal palmette) to be reinvigorated. Early versions of the free
(Fig. 15.9 and Plate 15.10). The most common palmette also utilized semi-vigorous root-
rootstocks used with this system are stocks, such as MM.106, but modern ver-
MM.106, MM.111 and M.7 (Rosati, 1978). sions utilize dwarfing rootstocks, such as
Tree spacing is 3.5–4 m in the row and 4–5 m M.9 or M.26, and higher tree densities.
between the rows (Corelli-Grappadelli, 2000). Common tree spacings are 1.5–2.5 m in the
Tree height is usually 4–5 m. The palmette row and 3–4 m between the rows. This gives
system has also been adapted to smaller a planting density of 1000 to 2200 trees ha1.
trees on M.9 with dimensions shown in Figs The trees are supported by a 3 m tall four- to
15.8 and 15.9. With the early versions of pal- six-wire trellis. Large, well-feathered trees
mette orchards, the tree was formed by are usually used. If feathered trees are used
annual heading cuts to the leader for the first to develop the palmette system, then the tree
several seasons, to obtain a very regular tree is planted without heading cuts (Table 15.7).
structure, with four to six tiers of either Some of the feathers that are not in the plane
oblique-angled or horizontal branches. This of the palmette (row plane) are removed and
the first tier of branches is selected from the
2.5 m remaining feathers. If the tree is not well
feathered, a heading cut is performed at
planting and the selection of the leader and
the branches must be delayed until the sec-
ond year. Heading cuts to the leader or lat-
eral branches are avoided to hasten the onset
of production. Summer pruning is used to
correct the growth of the tree. The tree is
allowed to grow freely in the first year after
planting. If it grows fast and develops suffi-
ciently, the second tier of branches can be
2.5 m selected during the first summer, or else the
Fig. 15.9. The horizontal-palmette tree has a
choice is postponed to the second growing
central trunk with four or five tiers of horizontal season. Higher-tier branches are developed
scaffold branches trained along the wires down the along the developing leader by growth regu-
row. A hierarchy of branch diameter is maintained lators or bending and twisting instead of cut-
from the bottom to the top of the tree by removing ting. They are selected during the second
large-diameter branches in the top. and third growing seasons, depending on
Table 15.7. Pruning and training plan for the palmette-trellis system. (NB. Northern-hemisphere dates.)
First year
using a flush cut. If tree has no feathers, head leader at 60 cm or expected height of
first wire. If the tree has two good feathers originating near expected height of the
bottom wire, head at 110 cm above soil line or at expected height of second wire.
Do not head scaffold branches
Early summer Install trellis support system with bottom wire at 60 cm and support posts every 15 m
along row. Attach tree to bottom wire with a permanent tree tie, leaving a 5 cm
July Tie developing leader to second wire with permanent tie. Remove clothes-pegs.
Choose two scaffolds on opposite sides of the trunk and tie to the bottom wire.
Ensure that shoot tips remain at a slight upward angle
Second year
Dormant If tree was headed to second wire in first year, skip to third-year training, otherwise
head leader at 5 cm above second wire
the leader shoot
July Tie developing leader to third wire with permanent tie. Remove clothes-pegs. Tie
bottom-tier scaffolds flat and attach to first wire with permanent ties. Choose two
limbs on opposite sides of trunk arising at second wire and tie to second wire
Third year
Dormant Head leader 10 cm above third wire. Remove unwanted, vigorous, upright shoots
along the bottom scaffolds. Remove vigorous upright limbs that are competing with
the leader
the leader shoot
July Tie developing leader to third wire with permanent tie. Remove clothes-pegs. Tie
second-tier scaffolds flat and attach to second wire with permanent ties. Choose two
limbs on opposite sides of trunk arising at third wire and tie to third wire
Fourth year
Dormant Head leader 10 cm above fourth wire. Remove unwanted, vigorous, upright shoots
along the bottom scaffolds. Remove vigorous upright shoots between tiers where
necessary
the leader shoot
July Tie developing leader to fourth wire with permanent tie. Remove clothes-pegs. Tie
third-tier scaffolds flat and attach to third wire with permanent ties. Choose two limbs
on opposite sides of trunk arising at fourth wire and tie to fourth wire
August Lightly summer-prune scaffolds on bottom two wires

Dormant Do not head the leader. Tie leader flat to the top wire in the fifth year and tie a vigorous
shoot in the opposite direction in the sixth year. Remove vigorous, upright shoots.
Remove upper-tier branches extending into the row, creating a flat fan arrangement
above the first wire. Shorten bottom-tier scaffolds where needed back to side-branch to
facilitate movement of equipment and preserve fruit quality on lower limbs
Continued
368 T.L. Robinson
Fifth and sixth year—Continued

July Tie fourth-tier scaffolds flat and attach to fourth wire with permanent ties
August Lightly summer-prune permanent scaffold branches by removing upright shoots
Mature-tree pruning
Dormant Limit tree to desired height by cutting leader back to top wire. Shorten productive
secondary branching on the bottom scaffold that extend into the alleys to facilitate
movement of equipment and preserve fruit quality on lower limbs. Shorten branches
on upper-tier scaffolds that extend out into the row to maintain narrow fan-shaped
top. Annually remove and renew one of the largest-diameter limbs on the upper
wires by tying a replacement shoot in its place
August Summer-prune by removing vigorous, upright shoots throughout the canopy. Be sure
to leave appropriate replacement shoots
the speed of development of the tree. The to be widely adopted in plums, sweet cherry
branches for each tier are selected to be and apricot (for the more vigorous cultivars).
80–100 cm from the next lowest tier without However, it is losing its edge to more inten-
complying to any geometric scheme. Selecting sive systems in apple and pear. In these
the branches in the summer has the advan- crops, where growth can be more easily con-
tage that they are easier to work with and trolled by dwarfing rootstocks, the palmette
will generally result in wider crotch angles. is giving way to higher-density systems
Selected branches are tied to the trellis, in the based on the spindle concept and its deriva-
plane of the row direction. Often the upper tions (e.g. super spindle) or on Y- or V-
branches are not permanent and are short- shaped canopies (Sansavini, 1993; Sansavini
ended to allow light penetration and can be and Corelli-Grappadelli, 1997).
renewed every 3–4 years.
Mature-tree pruning consists of removing
branches that extend across the row and 15.4.3 Penn State thin-wall trellis system
keeping the canopy confined to a 2 m wide
plane along the row. Upper-tier branches are The Penn State thin-wall trellis was devel-
renewed by removing one or two branches oped by Tukey (1978) and is a modification
per year and allowing replacement branches of the regular palmette system. It uses a low
to develop. Pruning the palmette is as quick 1.8 m tall four-wire trellis to allow all man-
as other systems (Sansavini et al., 1980). If the agement operations to be done from the
canopy is kept narrow, with good light pene- ground. It was also designed to be harvested
tration, and the trellis helps maintain the mechanically and a prototype over-the-row
shape throughout the life of the orchard, the harvester was built, which would comb the
tree may require less summer pruning than fruit off the trees. The system utilizes trees
other systems (Corelli and Sansavini, 1989). on dwarfing rootstocks, such as M.9 or M.26,
However, the pruning of the palmette system spaced at 1.8–2.0 m within rows and 3.0 m
cannot be neglected, as it too will develop between rows. The trellis has wires spaced
excessive growth, particularly at the top, with 45 cm apart. At planting, the tree is headed
all the disadvantages caused by shading. near the lowest wire and, from the branches
The palmette system in its various itera- that develop below the heading cut, two
tions has enjoyed broad success in Italy and scaffold branches are selected and trained in
southern France, where it still holds an each direction along the row at an oblique
important place in the cultivation of many angle (30°). The leader is headed annually at
crops and is still the training system of the next highest wire for the first 3 years and
choice for many growers. It is a leading sys- a pair of scaffolds are selected and trained at
tem for peach in the Italian Romagna region an oblique angle along the row. When the
(Correlli-Grappadelli, 2000), and continues leader reaches the top wire, it is bent along
the row and tied to the top wire to become a dwarfing rootstocks with closer in-row spac-
horizontal scaffold branch on the top wire. A ings. This canopy system was designed to
shoot arising from just below the top wire is match the tree form to mechanical harvesters
trained in the opposite direction to form a (Dunn and Stolp, 1987). A prototype mechan-
second scaffold on the top wire. Each of the ical harvester was built, which contacted the
scaffold branches is allowed to grow until it underside of the trellis, slightly raising it and
reaches the trunk of the next tree in the row. allowing the fruit to fall to a catching and
The scaffold branches cross at the mid-point conveying surface below. The tree is devel-
between trees, thus forming a permanent oped by planting a large caliper and tall
lattice of branches. The canopy develops into whip, heading the tree at 1.4 m and allowing
a continuous hedgerow. Mature-tree pruning four equal-diameter shoots to grow. Near the
consists of renewing the fruiting laterals that end of the first season, the four shoots are
arise off the scaffold branches by cutting trained horizontally along the row (two in
them back to the scaffold branch when they each direction) in the form of an H. These
become too long. The width of the trellis four shoots form the scaffold structure of the
hedgerow is determined by how long fruit- system. When vertical shoots that arise from
ing laterals are allowed to remain. the scaffolds are about 80 cm long, they are
The Penn State thin-wall trellis has had tied down to a horizontal position, perpen-
limited commercial acceptance in the north- dicular to the row direction. These shoots
east of North America. It has been very become the fruiting laterals of the canopy.
successful with difficult-to-colour cultivars, The canopy system creates a very regimented
such as ‘McIntosh’, where it has given the and regular, single layer of fruiting laterals at
best fruit colour. However, the low mature- 1.5 m high. By removing buds on the upper
tree height has resulted in relatively low side of the laterals, most of the fruits are pro-
yields. With vigorous cultivars or soils, duced on down- or side-buds. This allows
growers have had difficulty in controlling the fruit to hang free, which aids in mechani-
vigour in the tops of the trees as they age. cal harvesting.
The top horizontal scaffold branches have Only a few commercial plantings of
become excessively vigorous, producing many Lincoln canopy have been made. The hori-
strong vertical shoots each year, which causes zontal position of all of the branches of the
shading of the lower canopy. Although there tree has the inherent physiological weakness
have been some problems with this system, of stimulating excessive shoot growth from
it does have high labour efficiency for prun- the top of the canopy. This results in shading
ing and hand-harvesting. It continues to find of the fruit and spurs within the canopy
a place on pick-your-own farms, where the (Ferree et al., 1989) and a diversion of signifi-
trellis can also be used to keep people cant biological resources to the production of
picking in assigned rows. unwanted shoot growth rather than fruit.
Mechanical pruning was suggested by the
inventors of the Lincoln canopy system to
15.4.4 Lincoln canopy system remove the crop of vigorous shoots from the
top of the canopy. However, the indiscrimi-
The Lincoln canopy is a horizontal planar nate cutting of a mechanical saw exacerbates
canopy system developed by Dunn and Stolp this problem. The use of dwarfing rootstocks,
(1981) in New Zealand. The trees have a ver- such as M.9, would result in a lesser problem
tical trunk and a horizontal canopy at 1.5 m than the use of semi-vigorous rootstocks,
height (Plate 15.11). The width of the canopy such as MM.106.
is 1.2 m on each side of the tree row. Trees are
spaced 2.4 m in the row and 4.25 m between
rows. A 1 m gap between canopies of adja- 15.4.5 Ebro trellis system
cent rows is maintained. The system was
developed with semi-vigorous rootstocks, The Ebro trellis is a multi-tiered horizontal
such as MM.106, but could be adapted to planar system developed by a New Zealand
370 T.L. Robinson
commercial fruit-growing company in the growth from the top of the upper tier has
late 1970s (Tustin et al., 1989). The system been difficult to control. To improve the
utilizes an elaborate trellis structure with light-distribution characteristics of the Ebro
four horizontal tiers of wires stacked on top trellis, several modifications were proposed:
of each other and spaced 0.5 m apart, with (i) widen the bottom layer and narrow the
the lowest tier at 0.9 m high and the top tier top layer to give a triangular cross-sectional
at 2.4 m. Each tier of wires has six strands of view of the canopy; (ii) reduce the number of
wires (three on each side of the tree) spaced layers to three instead of four; and (iii) use
25 cm apart, giving a total width of the trel- dwarfing rootstocks and closer in-row spac-
lis of 1.5 m. Trees are planted at 2.4 m in the ing to reduce tree vigour and the amount of
row and 3.6 m between rows. The trees are unwanted shoot growth. A grower in Long
trained with one or two vertical leaders up Island, New York, has been very successful
between the centre wires, with horizontal with a modified Ebro trellis that incor-
tiers of branches originating at each tier of porated all three design modifications.
wires. The original system utilized trees on
semi-vigorous rootstocks, such as MM.106,
but it has been adapted to trees on dwarfing 15.4.6 Solen system
stocks, such as M.9, in some parts of the
world. The canopy is developed by planting The Solen system was developed by
large caliper and tall whip trees, which are Lespinasse (1989) for use with tip-bearing
headed at the height of the first tier of wires (Type IV) cultivars. The Solen system is a
at planting. A new leader and four lateral low-domed (umbrella-shaped) form of
shoots of equal diameter are allowed to training. The systems utilizes dwarfing
grow. Near the end of the first season, the rootstocks, such as M.9. Trees are spaced
four lateral shoots are trained horizontally 1.5 m in the row and 4.5 m between rows.
along the row (two in each direction) in the Mature tree height is 2.0 m. The trees are
form of an H in a manner similar to the supported by a short two-wire trellis (wires
Lincoln canopy system. When vertical at 1 m and 1.6 m). The trees are developed
shoots that arise from the four scaffolds at by heading the tree at planting at 1.2 m
each tier are about 30 cm long, they are tied above the ground. Two main scaffold
down to a horizontal position, perpendicu- branches are developed below the heading
lar to the row direction. These shoots cut and, late in the first year or in the mid-
become the fruiting laterals of the canopy. In dle of the second season, are bent down the
the second, third and fourth seasons, the row to the opposite direction from where
process of heading the leader, developing they originate and tied to the top wire. The
four scaffolds and then fruiting laterals is two scaffolds thus cross each other at a
repeated at each additional tier. Once the point over the tree trunk. Fruiting laterals
canopy is mature, the fruiting branches are are developed off the two scaffolds. With
maintained by annual removal of the verti- tip-bearing cultivars, these fruiting
cal shoots arising from the top side of the branches bend readily with the weight of
horizontal branches. The objective is to fruit, forming a cascading wall of fruiting
develop a compact, efficient, high-yielding branches on each side of the trellis. Mature
orchard system that is adapted to the use of pruning consists of thinning out fruiting
mechanical production aids. branches to open up the canopy to maintain
Several hundred hectares of Ebro trellis good light exposure to all parts of the
orchard have been planted around the canopy and to renew the fruiting laterals.
world. However, as the trees have matured Although early results with this system
the chief challenge has been maintaining showed high yields, most of the commercial
adequate light exposure for the lower tiers of plantings had lower yields than taller sys-
the trellis to maintain fruit colour and consis- tems, such as the vertical axis. This system
tent cropping (Tustin et al., 1989; Warrington has limited commercial interest and only for
et al., 1996). In addition, vigorous shoot tip-bearing cultivars.
15.4.7 Tabletop bed system A second advantage of planar canopies is

that the thin two-dimensional canopy has
The tabletop bed system was developed by good light distribution to all parts of the
Preston (1978) and Palmer and Jackson (1977) canopy. The planar vertical trellis systems
at East Malling Research Station in England. It (palmettes and Penn State trellis) have pro-
is a very high-density system that eliminates duced some of the best fruit quality of any
the tractor alleys from between the tree rows, system. However, the horizontal planar sys-
leaving only a walking path. The beds can be tems (Lincoln and Ebro) have often had the
from three to 14 rows wide with a tractor poorest fruit colour (Ferree et al., 1989). The
alley between beds. This system utilized the Lincoln canopy and the original Ebro trellis
very dwarfing rootstock M.27. Spacing was proved to have significant management
0.5–1.0 m between trees and 1.5 m between problems, resulting in poor fruit quality,
rows. Mature tree height was 1.5 m and the and were abandoned. Both systems had
tree canopy resembled a table top. The trees excessive shoot growth arising from the tops
were supported by a short trellis only 75 cm of the horizontal canopies, especially when
tall. The trees were developed by annually semi-vigorous rootstocks were used.
shortening the leader and 1-year laterals to Another advantage of planar trellis sys-
one half of their length for the first 3 years. tems is that the tree training can be system-
Because of the very weak growth habit with atized. Training recipes can be developed
M.27 rootstocks, this style of pruning resulted where limb or branch training can be simpli-
in a shallow canopy of leaves and fruit fied and performed by unskilled labour. The
around the central leader. After the third year, trellis can serve as a template of tree shape.
pruning consisted of thinning out branches to The primary disadvantage of flat planar
maintain good light exposure to all parts of systems is that the labour costs to manipu-
the canopy. This system achieved very high late the canopy into a confined geometric
production (55 t ha1) and light interception shape are much higher than systems that are
(80%), with good fruit quality by the second based on a more natural tree shape.
season (Palmer, 1988). Mature yields were Attempts to reduce tree-training costs have
very high, due to the high light interception. partially succeeded, as with the free palmette
However, the very high cost of establishment system. A second and more critical disadvan-
of the system and the necessity of specialized tage is that most flat planar systems use rela-
equipment to control pests and manage har- tively low tree densities, which limits early
vest have limited the interest in this bed sys- yield and profitability.
tem. It has not been planted commercially.
15.5 V-shaped Canopy Systems

15.4.8 Present status of flat planar systems
Although V-shaped apple canopies have
The primary advantages of flat planar sys- existed for centuries in European gardens, it
tems are the ability to mechanize orchard was not until the 1970s that V-shaped fruit
management and picking operations. The pal- trees were popularized for commercial
mette was widely adopted in Italy and France orchards by Chalmers and van den Ende
because of the improved labour efficiency (1975) with the growing system known as
provided by picking platforms. Despite the the Tatura trellis. The Tatura trellis system
potential of completely mechanized harvest was originally developed for peaches to
with two-dimensional systems, this goal has increase yields and to allow mechanical
not been realized. Several prototype har- harvesting but was later adapted for apples
vesters were built, but their cost and their (Chalmers et al., 1978). A number of varia-
small but significant fruit-damage level, cou- tions of the Tatura trellis have been devel-
pled with the continuing adequate supplies of oped. Most of the newer V systems now
labour in most apple-producing areas in the utilize dwarfing rootstocks. With all V
world, have prevented their implementation. systems, the objective has been to improve
372 T.L. Robinson
yield by improving light interception and to that dwarfing rootstocks, such as M.9 and
improve fruit quality by improving light M.26, are used and a much smaller trellis is
penetration to the centre of the canopy. needed (2 m high with four wires per side).
V-shaped orchard systems can be catego- The trees are planted at 1 m in-row spacings
rized by tree shape and branch-training pro- and 4 m between rows. Trees are trained in a
tocol. There are two basic shapes of inclined manner similar to the Tatura trellis by head-
canopies: Y-shaped trees, which have a ing the tree at 50 cm height at planting and
vertical trunk and two opposing arms of the allowing only two shoots to develop, with
tree trained to either side of the trellis, and one trained to each side of the trellis.
V-shaped trees, where the whole tree is Secondary branches are trained at about 45°
leaned to one side of the trellis while the next or, in some cases, 90° from the leaders (hori-
tree in the row is leaned to the other side of zontal along the wires) and are attached to
the V trellis. Branch-training recipes of V the wires.
systems can be categorized into two types:
palmette and spindle. With the palmette type
of branch training, the secondary branches 15.5.3 Geneva Y-trellis system
are trained either flat along the wires or at an
oblique angle across the wires, in both cases The Geneva Y trellis was developed by
giving a flat two-dimensional fruiting plane Lakso (Robinson et al., 1989) and is a Y-
for each arm. With the spindle type of shaped system with the arms of the trellis at
branch training, the secondary branches are 60° above the horizontal (Plate 15.12). In con-
not trained to the wires and a three-dimen- trast to the Tatura trellis, trees in this system
sional cylinder or cone-shaped canopy is have multiple scaffold branches on each side
maintained around the main trunk. of the trellis, trained in a fan-shaped arrange-
ment (Fig. 15.10). Dwarfing rootstocks, such
as M.9, M.26 or M.9 interstems, are usually
15.5.1 Tatura trellis system used. The trellis is 2 m high, with only three
wires per side (Fig. 15.10). The trees are typi-
The Tatura trellis was developed by cally planted at 1.5–1.8 m in-row spacings
Chalmers and van den Ende (1975) and is a and with 4 m between rows. At planting,
Y-shaped system with the arms of the trellis trees are headed at 70 cm above the ground
at 60° above the horizontal. Each tree has and six to ten side-shoots are developed
two main scaffold arms, and secondary (Table 15.8). Half of the branches are trained
branches are trained as a palmette. Semi- to each side of the trellis in a fan arrange-
vigorous stocks, such as M.7, MM.106 and ment. Secondary branches are tied to the
MM.111, are usually used. The trellis is 3 m wires and are trained between the main scaf-
high, with six wires per side. The trees are folds to fill in the fan on each side.
planted at close in-row spacings (1 m), The optimum angle for the arms of the
typically with 5 m between rows. Trees are Geneva Y trellis was studied by Robinson
trained by heading the tree at 50 cm height at (1992a, 2000a). Robinson found a broad opti-
planting. Only two shoots are allowed to mum angle between 50° and 70° above the
develop, with one trained to each side of the horizontal where yield was maximized when
trellis. Secondary branches are trained at an between-row spacings were wide. However,
oblique angle (45°) with respect to the main at the close in-row spacing of 3 m, the opti-
scaffold branch and are attached to the wires. mum angle was more vertical (between 65°
and 70°). The optimum angle for fruit size
was also around 60°, while fruit colour was
15.5.2 Mini-Tatura trellis system best on the most vertical angles. The best
yield efficiency was at intermediate angles of
The mini-Tatura trellis was developed by around 60°. An angle of 65–70° above the
van den Ende and is similar to the Tatura horizontal resulted in the best balance of
trellis in shape and branch training, except vegetative growth, cropping and fruit quality.
points to the top wires. Scaffold branches

are trained along the sticks. Dwarfing root-
stocks, such as M.9 and M.26, are used. The
2.5 m
trees are typically planted at in-row spac-
ings of 1.8 m for Mikado and 1.35 m for
Drilling and 3.5 m between rows. At plant-
2m ing, trees are headed at 70 cm above the
cm
60 ground and either three or four side-shoots
are developed. Each scaffold branch is
cm
60 trained like the trunk of a slender spindle,
60∞
with secondary branches arranged around
60
cm the scaffold in a conic shape. The secondary
branches are not tied to the wires.
cm 15.5.5 MIA trellis system (A-shaped trellis)

75
The MIA trellis was developed by Hutton et

al. (1987) and is a double-row V-shaped
system, where arms of the adjacent rows of
the trellis form an A-shaped canopy over
2.25 m the tractor alleyway (Plate 15.13). Workers
Fig. 15.10. The Geneva Y-trellis tree consists of a have access to the centre of the V between
divided canopy with four or five equal-diameter the double rows. In many respects, it is a
branches trained to each side of the Y and no central modification of the Tatura trellis, but the
trunk. Scaffold branches are attached to the wires of adjacent double rows are leaned over to the
the trellis and spaced equally along the trellis. Large- trellis, creating an A shape under which
diameter scaffolds are removed and replaced. equipment travels. The double rows are
planted 2 m apart, while the inter-row space
beneath the canopy arms of the A shape is
15.5.4 Mikado and Drilling systems 6 m. The trees are planted at close in-row
spacings of 1–1.5 m. Trellis arms are 60°
The Mikado and Drilling systems were above the horizontal and the trellis is 3 m
developed by Widmer and Krebs (1997) and high, with six wires per side. Semi-vigorous
are Y-shaped systems, with the arms of the stocks, such as M.7, MM.106 and MM.111,
trellis at 70° above the horizontal. In the are usually used. Since the whole tree is
Mikado system, each tree has four scaffold inclined over to the angled trellis at plant-
branches of equal vigour (two on each side ing, large feathered trees are ideal and
of the trellis) originating from a single require very little pruning at planting. The
trunk. Each scaffold branch is trained to a main trunk is trained up the angled trellis
different quadrant of the trellis. With the and secondary branches are trained at an
Drilling system, each tree has three scaffold oblique angle (45°) from the trunk and are
branches, with two trained to one side of attached to the wires in a manner similar to
the trellis and one to the other. This the Tatura trellis.
arrangement is alternated down the row.
The trellis for both systems is 2 m high,
with only one top wire per side and a single 15.5.6 Mini-V-trellis system
low wire at 30 cm above the ground run-
ning alongside the trunk. Four bamboo The mini-V trellis was developed by Tom
sticks for Mikado and three sticks for Auvil of Washington State and is a V-shaped
Drilling are attached near the trunk to the system that is similar to the mini-Tatura
low wire and then at separate quadrant trellis, except that whole trees are leaned
374 T.L. Robinson
Table 15.8. Pruning and training plan for the Geneva Y-trellis system. (NB. Northern-hemisphere dates.)
First year
using a flush cut. Head leader to 60 cm or at the height of the expected base of
Y-trellis frame
1–2 cm growth Rub out the leader bud and attach clothes-pegs to ALL other new shoots to develop
four to six equal-vigour scaffold branches. Deflower tree
Early summer Install tree-support system and string a low training wire. Attach tree to training wire
but leave scaffold shoots in an upright untrained position
July Remove clothes-pegs
Second year
Dormant If fewer than six branches per tree developed in year 1, head sufficient scaffolds by
removing two-thirds their length to produce a total of ten scaffold limbs. Assume
three scaffolds will be created at each heading cut. DO NOT HEAD SCAFFOLDS if
there are six or more suitable branches per tree. Remove overly dominant branches
Mid-July Divide scaffolds and train one-half of them to each side of the Y, using a Max
Tapener. Uniformly space scaffolds on each side of the trellis in a fan arrangement. If
tips of trained branches are not upright, use string or rubber bands to tie in an
upright position to ensure continued extension growth
Third year
Dormant DO NOT HEAD SCAFFOLDS. Remove shoots and suckers that are too low and not
suitable for training to the wire
Mid-July Attach scaffold limbs to the second and third wires, maintaining tips of branches in
an upright position. Use permanent plastic trellis clips to attach scaffolds to first and
second wires, making sure to maintain fan arrangement. Train new shoots to the
trellis to fill existing gaps in the canopy. Remove unwanted upright sucker growth
Fourth year
Dormant Do not head scaffold limbs. Remove any low shoots or suckers that are not suitable
for training to the trellis wires. Minimize pruning to encourage cropping
August Attach scaffold branches to top wire, using trellis clips where necessary. Train
suckers that will be used as replacement shoots to the bottom wire. Remove
unwanted interior sucker growth
Fifth year
Dormant Minimize pruning. Where scaffold length is excessive, shorten scaffold length by
cutting back to side-branches just above the top wire
August Summer-prune by removing unwanted interior sucker growth. Retain one shoot on
each side and train to the bottom wire to be used as replacement scaffolds
Mature-tree pruning
Dormant Begin limb renewal pruning by removing one or two scaffold limbs each year.
Candidates for removal are those scaffolds that cause excessive crowding or are
excessively long. Maintain a minimum 1.5 m open gap between scaffold tips of
adjacent rows by the limb-renewal process and by shortening back long scaffolds to
a suitable side-branch above the top wire
August Summer-prune by removing unwanted interior sucker growth. Retain one shoot on
each side and train to the bottom wire to be used as replacement scaffolds. Maintain
a 1.5 m wide gap between rows by cutting back to suitable side-branches above the
top wire
alternately down the row to each side of the 15.5.8 V super-spindle system
trellis. The central leader of each tree is
trained up one side of the trellis. Dwarfing The V super spindle is a very high-density
rootstocks, such as M.9 and M.26, are used version of the V slender-spindle system.
and trees are planted at 1 m in-row spacings Trees are planted at 0.5 m in-row spacings
and 4 m between rows. The trellis is identical and 3.0 m between rows. The trees are leaned
to the mini-Tatura trellis. Since the whole tree alternately to each side of the trellis, as with
is leaned to the trellis at planting, large feath- the V slender spindle (Plate 15.15). The trellis
ered trees are ideal and require very little is also similar to the V slender spindle.
pruning at planting. The main trunk is trained Dwarfing rootstocks, such as M.9 and M.27,
up the angled trellis and secondary branches are used. Because of the very close in-row
are trained at an oblique angle (45°) from the spacings, individual trees are trained simi-
leader or in some cases at 90° from the leaders larly to the super-spindle system developed
(i.e. horizontal along the wires) and are by Fritz Nüberlin. The ideal tree has medium
attached to the wires. calliper, with many short feathers, which are
not pruned at planting. The main trunk is
trained up the angled pole without leader
15.5.7 Gütingen V slender-spindle system bending, while secondary branches are not
tied to the wires. Any large secondary
The Gütingen V was developed by Krebs branches that develop are removed and small
(1988) and is a V-shaped system, with indi- secondary branches are allowed to crop and
vidual conic-shaped trees, that allows high bend down. There are no permanent scaffold
tree densities without multiple tree rows branches in the tree.
(Mantinger, 2000). The trees are leaned alter-
nately to each side of the trellis and are
2.5 m
trained like conic slender-spindle trees (Fig.
15.11 and Plate 15.14). The trellis has an
angle of 75° above the horizontal and has
only one wire per side at 2 m high (Fig.
15.11). A 2.5 m steel, bamboo or wooden 1.8 m
pole is placed in the ground beside the tree
trunk and then leaned out to the wire
(Hoying and Robinson, 1993). The trees are
trained up the angled poles. Dwarfing root-
40∞
stocks, such as M.9 and M.27, are used and
trees are planted at 0.9 m in-row spacings
and 3.5 m between rows. Since the whole
tree is leaned to the trellis at planting, large
feathered trees are ideal and require very lit- cm
75
tle pruning at planting (Table 15.9). The
main trunk is trained up the angled pole,
using leader bending in a manner similar to
a leaned slender-spindle tree. Secondary
branches are not tied to the wires. The lower 2m
tier of branches is tied flat with strings,
Fig. 15.11. The Gütingen V-slender-spindle tree
while upper branches are trained flat and consists of a slender-spindle-trained tree leaned 20°
kept shortened or renewed when too long. away from vertical and attached to a support pole.
This keeps the tree in a cone shape for good Alternating trees are leaned to opposite sides of the
light penetration. V trellis.
376 T.L. Robinson
Table 15.9. Pruning and training plan for the V-slender-spindle system. (NB. Northern-hemisphere
First year
At planting Plant highly feathered trees. Adjust graft union to 10 cm above soil level. Remove all
scaffolds below 60 cm, using a flush cut. Trees with three or more scaffolds (25 cm
long) should be headed at uppermost scaffold. The uppermost side-branch should
then be tied up as the new leader. Scaffolds longer than 45 cm should be headed by
removing one-third their length
Soon after Install V-shaped-trellis system with an individual angled tree stake at each tree. The
planting support system should allow trees to be supported to 2.5 m. Attach alternating trees
to each side of the V-support system by leaning the whole tree to the trellis. Attach
tree to support system with permanent ties, leaving a 5 cm diameter loop to allow for
trunk growth
5–10 cm growth Attach clothes-pegs to new side-shoots on leader to promote wide crotch
angles
Second year
Dormant Where growth has been good (> 45 cm of terminal growth), tie leader and vigorous
scaffold branches horizontal. Alternatively, remove leader down to a horizontal
scaffold branch and tie it up as the new leader
Mid-June Tie leader back to stake, allowing smoothly curving bends to remain. Hand-thin fruits
to 15 cm apart
August Tie up lower scaffolds not expected to support the crop
Third year
Dormant Where previous leader bending has significantly weakened the top of the tree, DO
NOT PRUNE. Where leader is vigorous, tie leader horizontal or alternatively remove
leader down to a suitable horizontal side-branch and tie it up as the new leader.
Remove upper branches that extend into the interior of the V
Mid-June Tie leader back to stake allowing smoothly curving bends to remain. Tie down
vigorous limbs that will not bend with the weight of the crop. Hand-thin fruits to
15 cm apart
but prune back scaffolds to prevent limb breakage and preserve tree structure.
Remove shoots in the interior of the V
Fourth year
Dormant DO NOT PRUNE LEADER. Remove upper branches that extend into the interior of
the V. Remove overly vigorous upright limbs that are more than two-thirds the
diameter of the leader, using a bevel cut. Tie down other vigorous, upright limbs
below the horizontal overlooked during third summer
Mid-June Tie leader to stake, allowing smoothly curving bends to remain. Tie up fruitful
scaffolds that will not support crop weight
shape. Remove shoots in the interior of the V. Tie up lower scaffolds not expected to
support the crop

Dormant Limit tree height by cutting leader into 2- or 3-year-old wood back to a horizontal
fruiting branch. Remove upper branches that extend into the interior of the V. In each
year, remove at least one undesirable bottom-tier scaffold until four remain. Shorten
bottom-tier scaffolds by pruning back to side-branch to facilitate equipment
movement and preserve fruit quality on lower limbs. Shorten branches that have
become pendant back to horizontal portion of the branch
shape. Remove shoots in the interior of the V
Mature-tree pruning
Dormant Limit height by cutting leader back to a fruitful side-branch. Remove upper branches
that extend into the interior of the V. Shorten bottom-tier scaffolds by pruning back to
side-branch to facilitate equipment movement and preserve fruit quality on lower
limbs. Shorten branches that have become pendant back to horizontal portion of the
branch. Remove one or two vigorous upper scaffold limbs each year, preserving all
weak-fruiting wood and permanent lower-tier scaffolds
August Summer-prune to encourage light penetration and maintain pyramidal tree shape.
Remove shoots in the interior of the V
15.5.9 Advantages and disadvantages of the tree canopy with V systems since the
V systems canopy grows up over the tractor alleyways.
Typical between-row spacings for the Tatura
The benefits of V systems have led to a large trellis are 6 m, for the Geneva Y trellis 4 m
increase in their popularity over the last 25 and for the V slender spindle 3.5 m. The
years. In some parts of the world, V systems ability to use existing equipment in new V-
account for a significant portion of new apple shaped orchards reduces the equipment cost
plantings. The primary advantage of V sys- when converting old, low-density orchards
tems is that they have very high yields at to new, modern, high-yielding orchards. From
maturity (Hutton et al., 1987; van den Ende a tree-canopy management and yield per-
et al., 1987; Robinson and Lakso, 1989; spective, the highly rectangular planting
Robinson et al., 1991a; Robinson, 1992a). The designs of the V systems direct most of the
high yields have been related to high levels of canopy growth across the row over the trac-
light interception (Robinson and Lakso, 1991). tor alleyway, whereas most other systems
V systems allow less light to fall on the tractor direct growth both across the row and down
alleys between rows as a result of the arms of the row into the next tree. Research with
the V-shaped canopy growing over the tractor V systems has shown that cumulative yield
alleyways. With pyramid-shaped trees, con- over 11 years increased with increasing
siderable light is lost between rows and simi- rectangularity, up to a rectangularity of 4
lar levels of light interception to V systems are (between-row spacing four times greater
only achieved with multiple-row systems or than in-row spacing) (Robinson, 1997b). In
with very tall trees (Robinson and Lakso, contrast, research with conic-shaped trees
1989). Yields of V-shaped systems are also has shown that square designs are better,
higher than those of other systems, primarily since the canopy of a conic-shaped tree
due to the higher tree densities utilized by the extends down the row and across the row in
V systems. Direct comparisons of the V shape equal distances. With conic-shaped trees, the
with other tree forms at the same tree density highest yields are with a rectangularity of 1
have shown that the V systems have consis- (Parry, 1978).
tently yielded more than other vertical sys- Improved light distribution within the
tems by 8–15% (Robinson, 1997a). canopy has been another presumed advan-
Growers have observed that another tage of V-shaped canopies. V-shaped canopies
advantage of V systems is that the rectangu- are usually two-dimensional and should
lar planting designs used in V systems, have better light exposure than spherical or
where in-row spacings are much smaller conic-shaped trees, which are three-dimen-
than between row spacings, allow them to sional. However, light distribution depends
be managed with conventional equipment, on light transmission through the canopy
whereas the bed or very high-density and, if the canopy is too dense, the underside
systems require very narrow equipment. In of the trellis can be heavily shaded
most cases, tractors can drive under part of (Robinson and Lakso, 1991). In addition, the
378 T.L. Robinson
underside of the trellis can be heavily shaded for a vertical-axis system at the same density.
if the canopy arms of adjacent rows grow A second disadvantage is the greater cost
together. Our experience with this system to train the trees over the first 4 years com-
indicates that a minimum of 1.5 m of open pared with the central-leader or vertical-axis
space between canopy arms of adjacent rows systems. We have estimated that the labour
is necessary for good fruit colour. cost for tree training in years 1–4 is about
Another advantage of V systems, like the US$5000 ha1, compared with US$3500 ha1
planar canopy systems, is that the tree train- for the vertical-axis system.
ing can be systematized. Training recipes can A third possible disadvantage is that fruit
be developed where limb or branch training size from V systems is smaller than from ver-
can be simplified and performed by un- tical systems. Robinson et al. (1991a) have
skilled labour. This is aided by the trellis, shown that with ‘Empire’, fruit size with the
which serves as a template of tree shape. Y trellis was smaller than with the central-
Another reported advantage of V systems leader system. However, the Y trellis consis-
is reduced fruit sunburn in high-light- tently had heavier crop loads than the
intensity climates. Good data showing central-leader system, which could explain
reduced sunburn are lacking. However, in an the smaller fruit size. Statistical adjustment
unreplicated trial, van den Ende et al. (1987) of fruit size for crop load showed no sig-
reported lower fruit sunburn on ‘Nijisseiki’ nificant difference between the systems.
Asian pears when grown on Tatura trellis Nevertheless, the shape of the V systems
than when grown using central-leader, leads to high light interception and hence
multiple-leader or bush-trained trees. More high heat loads at midday, which could lead
solid data on sunburn are needed before this to greater water stress on the V systems. This
advantage can be claimed. could result in smaller fruit size.
The possibility of mechanical harvesting Growers who have adopted a V system
is another advantage of V systems (Chalmers have generally done so because of the benefit
et al., 1978). Van Heek and Adem (1980) and of increased yield or the presumed benefit of
Gould et al. (1986) have reported on mechan- improved fruit quality (primarily reduced
ical harvesters for Tatura-trellis peaches. sunburn and improved red colour). The ulti-
Removal averaged 95%, with 5% lost to the mate value of V systems depends on their
ground. Damage was minimal with plums economic performance. Although they have
but greater with peaches. Robinson et al. higher yields than other systems, they also
(1990) showed that Y-trellis apple canopies have higher costs. V systems will probably
had less damage when mechanically har- be better than conic-shaped tree systems
vested with a trunk-impact shaker than under conditions of high sunburn and high
central-leader-trained trees. This was primarily winds or where all the fruit must be picked
the result of placing the catching pads near from the ground, since the V systems can
the fruit, thereby reducing the distance fruits intercept more light with short-stature systems
fall and the possibility of contacting other than pyramid-shaped systems.
fruits or branches. If hand-harvest labour
becomes limiting, the V-shaped canopies offer
the best potential for mechanical harvest. 15.6 Tree-support Systems
The primary disadvantage of V systems is
the high cost of establishment and initial tree Tree-support systems were originally
training. Trellis costs of V systems are gener- designed to prevent tree leaning with poorly
ally more expensive than the costs of conic anchored rootstocks, such as M.9 and M.26.
or vertical planar systems. The cost of an However, modern pruning and training
installed trellis for a V system is around schemes, combined with dwarfing rootstocks,
US$6500 ha1 and, when added to the cost of favour the production of fruit before the
trees (US$7500 for 1500 trees ha1), results in tree’s own canopy can support the crop.
an establishment cost of around US$14,000 Thus modern tree-support systems are
ha1. This compares with about US$12,000 designed to support much of the weight of
the crop to prevent the branches of the tree developed and were based on short-stature
from breaking. The preservation of the tree trees, the most common tree support was a
canopy during the early years often depends 2.5 m long, round wooden pole placed at
on the engineering of a support system. A each tree. In southern Europe and North
good support system for a high-density America, where taller trees were common,
orchard must be viewed as an investment trellises were the norm and utilized larger-
that allows fruit production in the early diameter 3.5 m long wooden poles to support
years while preserving the tree canopy for three to six wires. As growers have tended
future, large, mature yields. In reality, the towards higher and higher densities, the
support system of a modern orchard is an most common support system in the world
investment that provides large returns rather today is a hybrid trellis with inexpensive
than being just an expense. Where trees are individual tree stakes (Fig. 15.12). This sys-
properly supported, larger trees can be tem has a strong anchor system at each end
grown in the first 5 years, resulting in larger of the row, with wooden support poles
crops each year. Without a proper support spaced at 15 m along the row. A single wire
system, trees must be pruned more heavily is strung along the tops of the poles and
to stiffen the wood to support the crop. This tightened to a high tension. At each tree, an
results in smaller trees and smaller crops. inexpensive individual tree stake is installed
Trees can be supported either by indi- and tied to the trellis wire. Individual tree
vidual tree stakes or by a trellis. Historically, stakes have been made from wood, bamboo,
the conic-shaped canopy systems were sup- galvanized steel conduit pipe or recycled
ported with individual tree stakes, while the angle iron. The tree is trained up the support
planar and V-shaped canopy systems were pole until it reaches the wire, when it is tied
supported by a trellis. Currently, the choice to the wire. This system provides the lower-
of tree-support method is often an economic cost benefits of a trellis and the management
decision. Many growers prefer the use of flexibility of an individual tree stake.
individual tree stakes over a trellis, since it The materials used for tree support
allows workers to move around the tree dur- around the world depend on the materials
ing harvest. With low and moderate tree available in that region and the growers’
densities, it is often economic to provide ingenuity. Growers in western North
individual tree supports. However, with very America, northern Europe and New Zealand
high tree densities, it becomes uneconomic have had ready access to inexpensive chemi-
to use individual tree stakes and a trellis cally treated wooden posts that do not rot.
must be used. In northern Europe, where Such posts are not as readily available in other
high-density orchard systems were first regions at the same cost. In some locations,
Wire
tensioner 10 cm Round or
square post 10
cm cm
10
3m
s t 3 m Steel stake Po
10 cm Po st
60∞ 3m 60∞
60∞ 60∞ 15 m
2.5 m
1.25 m 75 cm
cm 1m
75
Fig. 15.12. The hybrid trellis with tree-stake support system for both conic and V-shaped canopy systems
with either bamboo or steel-pipe tree stakes.
380 T.L. Robinson
untreated black locust poles are commonly Support systems vary widely in cost
substituted, with equal results. In areas of (Table 15.10). The least expensive high-den-
the world where wood is not available or is sity support systems are the vertical three-
prohibitively expensive, growers have been to five-wire trellises, while the most expen-
successful using reinforced-concrete posts. sive systems are the slender spindle and the
Trees on dwarfing rootstocks, such as elaborate trellis systems, such as the V slen-
M.9, must be supported to the desired der spindle, Tatura, Ebro or Lincoln. The rela-
height of the mature canopy. With the slen- tively greater cost of using individual
der-spindle system or the Penn State thin- wooden poles for tree support with the slen-
wall trellis, the support system only needs der-spindle system has prompted many
to be 2 m high since the systems are based slender-spindle growers to switch to a three-
on all work being done from the ground. wire trellis or a hybrid single-wire trellis,
However, with the taller systems, such as with either a steel-pipe or a bamboo tree
vertical axis, slender pyramid or HYTEC, stake for a support system. The super-spin-
the trees must be supported to 3 m high. In dle system, which is the highest-density
areas where long poles are inexpensive, it is apple system, utilizes a simple trellis and
common to use 3.6 m tall poles, with the top thus has one of the least expensive support
of the pole and the top wire at 3 m above the systems per hectare, but the very high tree
ground. In eastern North America, 3.6 m densities make it the most expensive
wooden poles are almost double the price of orchard system to plant.
3 m long poles, so a less expensive alterna-
tive, which still provides support up to 3 m,
is to use the shorter 3 m poles with the top 15.7 Field Comparisons of Orchard
of the pole and the top wire at 2.4 m above Planting Systems
the ground. At each tree a 12 mm diameter,
3 m long galvanized-steel pipe is used as an A number of field comparisons of planting
individual tree stake. The steel pole is systems have been conducted in the last 30
inserted in the ground only 15 cm, thus pro- years. In most cases, the trials have com-
viding a rigid support for the tree above the pared complete systems (unique combina-
wire (Fig. 15.12). tions of tree density, tree form, rootstock and
Table 15.10. Support-system costs for high-density orchard systems.
Tree Material Labour Total

density costs costs cost
(trees (US$ (US$ (US$
System ha1) Support-system description ha1) ha1) ha1)
Central leader 500 Temporary wood trunk stake (2 m 5 cm) 998 349 1348
Mini-central leader 850 Steel-pole tree stake (3 m 5 cm) 1470 336 1806
Free palmette 950 Five-wire trellis (3 m high) 2396 1937 4333
Penn State trellis 1100 Three-wire trellis (2 m high) 2497 1559 4056
Vertical axis, 1500 Hybrid trellis (single-wire trellis 2.5 m high 3252 1604 4856
slender pyramid with 3 m steel-pole tree stakes)
or HYTEC
Vertical axis, 1500 Hybrid trellis (single-wire trellis 2.5 m high 2499 1604 4103
slender pyramid with 3 m bamboo-pole tree stakes)
or HYTEC
Geneva Y trellis 1500 Y trellis (2.2 m high) with three wires per side 3025 2653 5678
Slender spindle 2000 Wooden tree stake (2.5 m 5 cm) 5202 4161 9363
V slender spindle 3000 V trellis (2.2 m high) with one wire per side 5549 3053 8603
and with 3 m steel-pole tree stakes
Super-spindle 5000 Three-wire trellis (2.4 m high) 2049 1721 3770
training system), which have made it diffi- central-leader system. Yields per hectare
cult to determine the individual effects of were largely a function of the planting den-
tree density, training system or rootstock. sity. As the orchard matured, the Y trellis had
Almost all field comparisons of systems have the greatest yield, while the yields of the
shown that the higher-tree-density systems three conic-shaped systems continued to be a
have had higher yields than lower-density function of tree density. This anomaly was
systems, at least in the early years. due to the greater light interception at matu-
Ferree (1980) studied four systems: slender- rity of the Y-trellis system than that of any of
spindle/M.9, palmette trellis/M.9, mini- the other systems. This same yield ranking
central leader/interstem and large central among the four systems continued until the
leader/MM.106. He found that cumulative end of the trial at age 22 (Robinson, 2000b).
yield was related to planting density, with In the early years, there were few differences
the slender spindle having the highest yield. in colour, but the high-yielding systems had
However, the soils and climate in Ohio gave slightly smaller fruit size than the low-
relatively high vigour, which resulted in yielding systems, due to heavier crop loads.
poorer light levels within the canopy of the In the second decade of the orchard’s life, the
slender-spindle trees than the palmette or mini-central leader had the best colour,
mini-central leader. The latter two systems followed by the large central-leader and
had more open canopies and greater crop slender-spindle systems. The Y trellis had
loads. The palmette system had the highest poorer colour in years 11 and 12 as the tops
yield efficiency, indicating the best balance of adjacent rows grew together. After correc-
between vegetative growth and cropping. tive pruning and the application of renewal-
Blizzard et al. (1988) compared slender pruning strategies to that system, fruit colour
spindle/M.9, palmette trellis/M.9, Lincoln was similar to that of the other systems. Fruit
canopy/M.26 and central leader/MM.111. size tended to be smaller with the Y trellis
The slender-spindle system, with a density than with other systems. It appeared that
of 4000 trees ha1, had the highest yield and this was largely due to heavier crop loads;
the central leader, with a density of 270 trees however, there also appeared to be a consis-
ha1, had the lowest yield. The palmette and tent but small effect of system on fruit size. It
the Lincoln canopy were similar in yield and may be that the high interception of light
had the same density. At maturity, the pal- during the midday hours results in larger
mette system had the most open canopy and heat loads and water stress on the Y-trellis
the best fruit quality, while the Lincoln canopy than on the conic-shaped trees.
canopy and the slender spindle had the Lespinasse et al. (1992) compared seven
poorest fruit quality (Ferree et al., 1989). The systems (vertical axis, structured axis, Tatura,
latter two systems had more dense canopies pergola, palmette, MIA 15°, MIA 30°) on
and poorer light exposure to the lower part ‘Royal Gala’ trees grafted on M.26. Results
of the canopy. In this study, the tree vigour showed that the best cumulative yield and
of the soil and climate gave too much fruit size were obtained with the four train-
growth for the high-density slender-spindle ing systems vertical axis, structured axis,
system and, despite high yields, gave poor Tatura and palmette. The vertical-axis system
fruit quality. was characterized by the best fruit colour,
Robinson and Lakso (1989) compared indicating that this system optimized light
four orchard systems (slender spindle/M.9 distribution within the tree canopy.
at 1960 trees ha1, Y trellis/M.26 at 1283 trees In a comparison of slender spindle and
ha1, mini-central leader/M.9/MM.111 at super spindle, Weber (2000) has shown
961 trees ha1 and large central leader/M.7 greater cumulative yield with the super
at 450 trees ha1) in one of the few trials to spindle compared with the slender spindle,
be conducted for longer than 20 years. Yields but reduced fruit size and fruit colour.
during the early years were highest for the Wagenmakers et al. (1994) also showed that,
slender spindle, followed by the Y trellis and although total production increased with
mini-central leader, and lowest for the large higher tree densities, the proportion of well-
382 T.L. Robinson
coloured fruit decreased sharply when tree fourth–eighth years, the Y trellis had slightly
density exceeded 10,000 trees ha1. In addi- higher yields than the other systems, but the
tion, the very close spacings utilized in the differences were not large. After 8 years, the
super-spindle system have resulted in Y trellis had the greatest cumulative yield
greater growth in the upper part of the tree (8–15% more), while there were no significant
compared with the lower part of the tree, differences among the other three systems.
thus increasing the risk of losing production These results indicate that any one of the
in the lower canopy zones, due to light defi- modern training systems will produce excel-
ciency and less vegetative growth at the bot- lent results if planted at the same density
tom part of the tree (Weber, 2000). Thus and on the same rootstock.
orchard light management with the super Buler et al. (2001) compared the slender-
spindle is very important to ensure produc- spindle, HYTEC, Solen and Mikado systems
tivity (Wagenmakers, 1995). in Poland. They found that the slender-
Orchard trials with V-shaped canopies spindle and Mikado systems had higher
have shown them to be highly productive yield than the HYTEC or Solen systems. The
and highly efficient at converting light Mikado system gave the best fruit size and
energy into fruit. Among four systems, the light distribution in the canopy.
Geneva Y trellis/M.26 had the highest con-
version efficiency (6.5 g fruit MJ1 photosyn-
thetically active radiation (PAR)), followed in 15.8 Underlying Principles of High-
order by the slender-spindle/M.9, interstem density Orchard Planting Systems
and central-leader/M.7 systems (4.0 g fruit
MJ1 PAR) (Robinson and Lakso, 1991). The Although modern orchard planting systems
greater efficiency of V systems is probably vary in specific tree-training recipes, they are
due to the good exposure of the fruiting based on many of the same underlying prin-
spurs in the Y-trellis system. ciples (Palmer and Warrington, 2000). These
Widmer and Krebs (2001) compared include high tree density, precocious and
slender-spindle, V slender-spindle, Drilling and dwarfing rootstocks, high light interception
Mikado systems at several densities. With at maturity, good light distribution within
increasing tree density, yield per tree the canopy and a balance between vegetative
decreased, but yield per hectare increased. growth and cropping.
However, the increase in yield per hectare
was proportional to the increase in trees per
hectare. At the highest tree density, fruit size 15.8.1 Tree density
was reduced. The open-tree forms of the
Drilling and Mikado systems had lower yield Tree density is the single most important
in the early years, but similar yield and better factor affecting early yields of any orchard
fruit quality when the trees were mature. system. In the early years of an orchard’s life,
To separate the effect of planting density light interception by the canopy is low, which
from training system, Robinson (1997a) limits yield potential. A major objective dur-
compared four training systems (slender ing the first few years is to develop the
spindle, vertical axis, Y trellis and horizontal canopy as quickly as possible so that full
palmette), each at two tree densities, using canopy closure can be achieved. With tree
two cultivars with distinct growth habits densities below 500 trees ha1 this can take
(‘Jonagold’ and ‘Delicious’), but all on the 7–12 years. With densities between 1500 and
same rootstock (Mark). Over the first 8 years, 3000 trees ha1, which are the most common
the higher-density version of each system densities in modern high-density orchards
produced more than the low-density version. around the world, tree-canopy closure can be
However, despite the very different training achieved by the end of the third or fourth sea-
strategies imposed on the trees over the first son. With the very high tree densities of the
5 years there was no significant difference in super-spindle system (> 4000), canopy closure
third-year yield among the systems. In the can be achieved by the end of the first year.
Data from several studies show that, is increased an additional cumulative yield
during the early years, yields are related to of 150 kg ha1 was obtained for each addi-
tree density, with the highest tree density tional tree per hectare. This would be about
producing the highest cumulative yield (Fig. eight times the cost of the additional tree. At
15.13). The relationship of tree density and the higher tree densities, the gain in cumula-
cumulative yield is linear in the first 2–3 tive yield was very small, with a slope of 70
years, but by year 6 the relationship is curvi- kg per tree for ‘Jonagold’ and 20 kg for
linear (Fig. 15.13). Data from New York also ‘Empire’. This would be about 3.5 and 1
show a curvilinear relationship over the first times the cost of the additional tree for
11 years of an orchard’s life (Fig. 15.14). At ‘Jonagold’ and ‘Empire’, respectively. In
the lower end of the density continuum, the northern Italy, a comparison between a
relationship is almost linear, with a slope of super-spindle orchard with 6143 trees ha1
150 kg per tree indicating that as tree density and a string-tree orchard with 13,134 trees
600
6-Year cumulative yield (t ha–1)
500
400
300
200
100
‘Jonagold’
‘Elstar’
0
0 5000 10,000 15,000 20,000
Tree density ha–1
Fig. 15.13. Relationship of tree density and cumulative yield over the first 6 years of an apple orchard in
The Netherlands (from Balkhoven-Baart et al., 2000).
500
‘Jonagold’
11-Year cumulative yield (t ha–1)
‘Empire’
400
300
200
100
0 1000 2000 3000 4000
Trees density ha–1
Fig. 15.14. Relationship of tree density and cumulative yield over the first 11 years of an apple orchard on
M.26 trained to Y-trellis system in New York State, USA (‘Jonagold’ r2 = 0.95; ‘Empire’ r2 = 0.79).
384 T.L. Robinson
ha1 showed that, with a twofold increase of cumulative yields, profitability is only
tree density, only a 1.17-fold increase in yield slightly greater and, in some cases, less than
in year 2 was obtained and only a 1.09-fold with lower densities; moreover, the economic
increase in yield in year 4 with ‘Golden risk is increased significantly with very high-
Delicious’ (Österreicher, 1993). The relation- density orchards. It should be noted, how-
ship of planting density and cumulative ever, that the relationship of tree density and
yield over the life of an orchard is typical of early cumulative yield can be modified by a
the law of diminishing returns, which states number of factors, including initial tree qual-
that additional increases in an input factor ity and pruning severity, which would
(tree density) produces a smaller and smaller change the optimum economic density.
increase in an output factor (yield). At the Tree density, when properly calculated,
high end of this curvilinear relationship, considers the vigour of the cultivar, the
additional increases in tree density will not vigour of the rootstock and soil strength.
produce enough extra yield to pay for the With vigorous scion cultivars, growers
additional costs incurred to purchase and should use a more dwarfing stock and
plant the extra trees (Weber, 2001; Widmer greater planting distances. With weak scion
and Krebs, 2001). cultivars, a more vigorous rootstock should
The optimum tree density in any apple- be used and/or closer planting distances.
producing area is an economic question. The Despite some latitude in planting distances,
laws of economics dictate that the optimum growers should remember that, to obtain
density will be less than the density with the high early yields, high tree densities are
highest yield. In Europe, average planting essential. Thus in almost all cases, planting
densities increased up to about 1995 to distances should not exceed 2.5 m in the row
5500–6000 trees ha1, but in the last 6 years or 4.5 m between rows.
there has been a trend towards more moder-
ate planting densities, ranging between 2800
and 3800 trees ha1 (Weber, 2000). The reason 15.8.2 Rootstock
why more moderate planting densities are
favoured may be explained by rising tree Although high tree density is the single most
prices and lower returns for the fruit. It has important factor affecting yield in the early
become more difficult to justify the high years of an orchard’s life, dwarfing rootstocks
investment cost of the super-spindle system. are the foundation for any successful high-
Another reason for more moderate plant den- density planting system. The choice of root-
sities is the difficulty of managing excessive stock in combination with the choice of scion
vigour, especially of virus-free plant material cultivar defines the typical tree vigour and
of new cultivars, such as ‘Gala’ and final tree size, which dictate its planting den-
‘Braeburn’ (Weber, 2000, 2001). Many grow- sity. Rootstock has a dominant effect on tree
ers have not been successful in balancing the precocity (flowering and cropping in the early
vegetative and reproductive growth of a years) and productivity. In many ways, the
super-spindle orchard. Our latest economic choice of rootstock determines the potential
analyses under New York conditions have yield of a given cultivar and ultimate prof-
indicated that densities between 1000 and itability. The rootstock also determines tree
2500 trees ha1 are more profitable than size, which determines a system’s labour effi-
lower or higher densities. The results of our ciency. Other rootstock characteristics impor-
economic analysis may not be valid for other tant to consider in selecting the right rootstock
parts of the world, with different fruit, tree include tolerance to diseases, such as fire
and land prices. Nevertheless, for most fruit- blight, Phytophthora and crown gall, tolerance
growing areas of the world, it is our opinion to insects, such as woolly apple aphid (WAA),
that tree densities up to about 2500 trees ha1 and tolerance to abiotic stresses, such as
will result in greater profitability and moder- drought and/or water tolerance, and spring-
ate levels of risk. Above this density, it frost and winter-cold tolerance. Rootstock can
appears that, despite producing greater also have a direct effect on fruit size.
Most successful high-density plantings ing on light interception. Light interception

are planted with dwarfing rootstocks, such is largely a function of tree shape and
as M.9, M.26, B.9, G.16, CG.202 or O.3, or arrangement.
with interstem trees of M.9 and MM.111 Mathematical models of light interception
or MM.106. Although it is possible to have been constructed to estimate light
plant high-density orchards on semi-dwarf interception from different canopy shapes
rootstocks, such as M.7 or G.30, or some and planting configurations (Jackson, 1981;
semi-vigorous rootstocks, such as MM.111, Palmer, 1981). From this body of work, a few
MM.106 or M.793, their lack of precocity is main points are important for this discus-
a serious limitation to this approach. In sion. Orchard canopies, unlike those of other
addition, their inherent vigour makes man- crops, are of necessity discontinuous, due
agement of the mature high-density orchard to the alleyways maintained between rows
much more difficult. Dwarfing rootstocks for orchard machinery, which results in a
limit tree vegetative growth when trees are large proportion of the land area not being
mature, which results in less winter and covered by trees (Jackson, 1980). This results
summer pruning. in low values for leaf area index (LAI) and
Within the M.9 family of dwarfing root- total light interception for orchard canopies
stocks, there are significant differences in when compared with other crops. In addi-
vigour between clones. The weaker clones tion, leaf area is not randomly distributed
(M.9 NAKBT337 and M.9 Flueren56) are over the land area, as in annual crops, but is
especially useful with vigorous scion culti- clumped in trees, branches and spurs. Light
vars on virgin soil. The more vigorous clones interception is low when the orchard is
(M.9 Pajam 2, M.9 Nic29, M.9 EMLA) are planted and increases as the orchard devel-
much better when orchards are planted on ops in relation to total LAI. Light inter-
replanted soil or when weak scion cultivars ception in orchards can be raised by: (i)
are used. Although M.9 and M.26 are used increasing the density of foliage in the
around the world with great success in high- canopy; (ii) increasing the height of the trees
density plantings, they are both susceptible relative to the clear-alley width; or (iii)
to fire blight and WAA. In addition, M.26 is increasing the number of trees per hectare
very susceptible to Phytophthora root rot. The (Corelli and Sansavini, 1989). Because of the
new dwarfing rootstocks that are resistant to tractor alleys used for orchard management,
these problems, such as the Cornell Geneva light interception is more strongly influenced
series, should improve the worldwide per- by tree numbers per hectare and ratios of
formance of high-density orchards. tree height to clear alleys than by canopy
density.
Cain (1970) introduced the idea that
15.8.3 High light interception at orchard orchard light interception should be consid-
maturity ered over the lifetime of the orchard. He
found that the mean lifetime fraction of land
Mature yields of all apple orchards, regard- covered by tree canopy increased as tree size
less of planting density or pruning system, decreased and tree planting density
are related to total light interception (Palmer, increased. A meaningful variable of orchard
1989, 1997; Barritt et al., 1991; Robinson and performance would be mean lifetime frac-
Lakso, 1991; Wagenmakers, 1991; Robinson, tion of light intercepted, but few studies
1992b; Robinson et al., 1993). The theory of have calculated such an index.
light interception has been reviewed several Jackson (1970) has shown that the widely
times (Cain, 1972; Jackson and Palmer, spaced bush trees common in many older
1972; Jackson, 1980, 1985; Sansavini, 1982). orchards, which are large at maturity, inter-
Although early yields appear to be largely a cept very little light when they are young
function of tree density and rootstock and and yet when full-sized they have excessive
not greatly influenced by training system, within-tree shading and relatively low LAIs.
mature yields can differ substantially depend- The realization that significant land and light
386 T.L. Robinson
resources were wasted in the early life of an spaced 1.5 m by 0.5 m in a 14-row bed
orchard has encouraged the planting of separated by tractor alleyways. It achieved
higher and higher tree densities. This has ceiling levels of LAI and light interception
resulted in greater early yield and greater 2 years after planting. Interception was com-
lifetime light interception. Verheij and parable with that of a closely planted
Werwer (1973) examined light interception in hedgerow system (2.9 m by 0.9 m) and yet
low- and high-density hedgerows of ‘Golden the bed system had a much lower LAI, due to
Delicious’. The low-density plots (1100 trees a more even distribution of foliage over the
ha1 on M.9 and 660 trees ha1 on M.2) inter- orchard floor. They concluded that LAI was
cepted roughly half of the incident light at the largest single factor influencing light
maturity, and yields peaked at 40 t ha1. The interception in discontinuous orchard
high-density plots (3300 trees ha1 on M.9 canopies; however, the LAI/light intercep-
and 2260 trees ha1 on M.2) intercepted 66% tion relationship could be modified to some
and 75% of available light, respectively. extent by leaf-distribution pattern over the
Yields were more than 70 t ha1 in their sixth ground area. More recently, Palmer (1988)
and seventh years, but thereafter yields has shown that bed systems can intercept up
declined due to inter-tree competition. In a to 80% of PAR from the end of June until
spacing trial, Palmer and Jackson (1974) have October and yield 78 t ha1 in the third year.
reported that, with densities ranging from The more complicated tree arrangements of
853 to 3746 trees ha1, yield and light inter- multi-rows and bed systems have been more
ception were approximately linearly related. difficult to manage than single rows.
Light interception was closely related to LAI, Intensive bed systems (Palmer and Jackson,
not just trees per hectare. 1977) or multi-row systems (Wertheim et al.,
Trees can be arranged as single rows, as 1986) are not compatible with conventional
multiple rows, as a bed system with occa- orchard machinery and have generally not
sional drive alleys or as meadow arrange- yet been adopted widely. Single-row arrange-
ments that have no drive alleys. The goal of ments continue to be the most successful.
each arrangement is to maximize the inter- Few studies of light interception have
ception of sunlight. Wertheim et al. (1986) been done with planar canopies. Palmer and
examined light interception of high-density Jackson (1977) reported that trees on semi-
single-row, three- and six-row bed and full- dwarfing rootstocks, trained to tall narrow
field systems. They found that light intercep- hedgerows separated by wide alleys,
tion was positively related to tree density achieved high light interception at maturity,
and to yield. The effect of the tree arrange- but the between-row shading resulted in
ment was less important than tree numbers poor illumination of the lower portion of the
per hectare for increasing total light intercep- hedgerow, which became unproductive.
tion and yield. The results of tree arrange- Shorter hedges on more dwarfing stocks
ment showed that a single-row or full-field inevitably have lower light interception if the
arrangement of trees gave better yields than alley width is maintained the same. Such
the three-row system at equivalent tree den- short, vertical, trellised hedgerows are com-
sities. Increasing tree planting density has mon in New York but, in many cases, tractor
been the most important means of increasing alley widths have remained too wide for
the early yield and early light interception of optimum light interception and yields have
young orchards. Barritt (1989) found that been lower than expected. The recent intro-
tree density was more important than train- duction of narrow orchard tractors to the
ing system or rootstock for improving light USA should help in reducing tractor alleys.
interception and yield in the third year with Robinson and Lakso (1989) studied light
‘Granny Smith’ apple trees. interception and yield among three conic-
Palmer and Jackson (1977) developed a shaped systems and a V-shaped system.
bed system to achieve high light interception Light interception was highest with the Y
with a more even distribution of foliage over trellis – 70% of available PAR – compared
the orchard floor. The bed system had trees with 55% for the slender-spindle system,
despite a higher tree density with the spacings used in high-density orchards of
slender-spindle system. The increased light 3.5–4.5 m should have mature tree heights of
interception was the result of the canopy 2.8–4.0 m. Failures of mature high-density
architecture, which allowed the tree canopy orchards to produce expected yields at matu-
to grow over the tractor alleys. There was a rity can usually be traced to trees not inter-
linear relationship between yield and tree cepting enough light. Where growers desire
density over the first 10 years of the orchard short-stature trees (< 2.5 m tall) that can be
life for the three conic-shaped systems, but picked from the ground, they should reduce
the Y-trellis system had greater yield than tractor alleys to 2 m or less.
was predicted for its tree density. The
increased light interception accounted for a
large portion of the increased yield. The Y 15.8.4 Good light distribution within the
trellis, because of its unique geometric shape, mature canopy
had interception levels similar to those of the
bed systems of Palmer (1988) or the multiple As trees mature, the challenge becomes to
rows of Wertheim et al. (1986) and yet could utilize canopy-management strategies that
be maintained with conventional equipment. maintain good light distribution within the
The light-interception properties of other pla- canopy. With all apple-planting systems,
nar-canopy tree forms have not been studied. there is a natural progression towards an
From the mid-1970s to the late 1980s, umbrella-shaped tree, where the upper limbs
there was a strong move toward small trees receive more light intensity and thus grow
that allowed all orchard management opera- more than the lower limbs, causing shading
tions to be done from the ground. In many of the lower branches. Shaded areas of the
cases, the decrease in tree size has not been canopy produce smaller fruit size, poorer
accompanied by a reduction in tractor alley fruit colour, less return bloom and weak
width and tree height was kept too low for fruiting spurs (Jackson, 1970; Jackson et al.,
high light interception. As a consequence, 1977; Robinson et al., 1983; Barritt et al., 1987;
many dwarf orchards have relatively low Lasko et al., 1989b; Palmer, 1989; Warrington
mature yields. Robinson and Lakso (1989) et al., 1996; Wunsche et al., 1996). Our work
found that a Y-trellis system intercepted with orchard systems has shown that there
about 70% of available PAR at maturity, are no important differences in fruit quality
while the slender-spindle system intercepted in the first few years of an orchard’s life.
only 55% of PAR, in spite of 30% greater tree However, as the orchards reach maturity,
density with this system. This illustrates the significant differences in fruit colour, size
problem of short-stature trees planted in and quality become apparent.
single rows, where mature light interception Two approaches have been used to
is relatively low due to a low ratio of tree improve the light distribution in mature
height to clear alleys. There is a relationship apple canopies. One is to provide and man-
between tree height, row spacing and light age many small openings between branches,
interception. Greater tree height at the same which allow light penetration to the centre
row spacing increases light interception and, and lower portions of the canopy. This
therefore yield, if light distribution to all approach is used with natural tree forms,
parts of the canopy is adequate (Jackson, such as in the central-leader, slender-spindle,
1980; Barritt et al., 1991; Robinson and Lakso, vertical-axis, slender-pyramid or HYTEC
1991; Robinson et al., 1991b; Wagenmakers systems (Wertheim, 1968; McKenzie, 1972;
and Callesen, 1995). For optimum light inter- Heinicke, 1975; Lespinasse and Delort, 1986;
ception, the ratio of tree height to row spac- Barritt, 2000; Tustin, 2000). This approach
ing should be 0.8–0.9. Many of the problem can be successful, but generally requires a
orchards in the 1980s, which had short trees high degree of horticultural skill to manage
and utilized existing wide machinery, had the growth of the canopy. A second approach
height-to-row spacing ratios of 0.5 or less. is to provide fewer, large, permanent
Using the optimum ratio, typical row openings for light penetration into canopies
388 T.L. Robinson
restricted into geometric forms, such as penetration depth of light into unrestricted
palmette hedgerows, tree walls and A, V or T apple canopies is about 1 m.
forms (Chalmers and van den Ende, 1975; Dense hedgerow trees were studied by
Luckwill, 1978; McKenzie et al., 1978; Rosati, Verheij and Werwer (1973), who found that
1978; Tukey, 1978; Dunn and Stolp, 1987; average light levels exceeding 50% of full
Hutton et al., 1987; van den Ende et al., 1987; sunlight occurred only at the top periphery
Palmer, 1988; Lakso et al., 1989a; Correlli- of the canopy. Moving down and inward in
Grappadelli, 2000). This approach generally the canopy, average light levels dropped
requires severe geometric restriction of the sharply to about 15% of full sunlight or less.
canopy, expensive support structures and The light penetration into large and small
significant labour to place and maintain the hedgerow trees was similar, but the cross-
branches in specific locations. The value of section of small trees allowed for better light
these different tree forms lies in their light- illumination of the interior portions of the
distribution properties and the attendant canopy (Verheij and Werwer, 1973). Heinicke
improvements in fruit yields and/or quality. (1964) also found that, as tree size decreased,
The work of Heinicke (1963) and Looney the heavily shaded area within the tree
(1968) showed that large, round-crown trees decreased. Leaf area per tree decreased with
have a large central core of the canopy that smaller tree size, but leaf area per hectare
receives very low light intensities (6–30% of increased. Dwarf trees with some overlap of
full sunlight). The exterior quarter of the canopies had one-third more leaf area per
canopy had a small percentage of the total hectare that received > 30% full sun than did
leaf area and yet had a large shading effect standard trees. This indicates a distinct
on the rest of the tree, where the major por- advantage in photosynthetic potential for
tion of the leaf surface was located (Fig. smaller trees. Forshey and McKee (1970)
15.15). Heinicke (1963) proposed that 30% of reported that a large and small tree had the
full sun serve as a lower limit of desired light same total dry-matter accumulation per unit
level in apple canopies. Jackson (1970) found of occupied land, despite the lower LAI on
a more rapid decline in light level with the smaller tree. The small tree had a more
depth of canopy, with light levels reduced to efficient leaf surface and produced 80% more
34% of full sun within 1 m of the canopy fruit per unit of occupied land than the big
exterior. He found that the main cropping tree. Cain (1970) showed a negative linear
zone of the tree received a minimum of 35% relationship between production per unit of
full sun, while the more shaded areas pro- tree area and the size of the tree. The rela-
duced relatively few fruits. This result has tionship showed a decrease of 0.6 kg m2 for
led to the rule of thumb that the effective each metre increase in tree spread. The
decreased efficiency of large trees is probably
the result of greater internal shading.
Robinson et al. (1991b) characterized the
71–100%
light climate of mature central-leader
51–70%
‘Empire’/M.7 and mini-central-leader
31–50% ‘Empire’/M.9/MM.111 trees (interstems),
slender-spindle/M.9 trees and Y-trellis M.26
trees at four times during the growing sea-
< 30% of son to examine the effect of tree size, shape
full sun and summer pruning on light exposure in
the lower part of the canopy. They catego-
rized the canopy into three light zones: (i)
well-exposed zone where light levels
exceeded 50% full sun; (ii) marginally
exposed zone with light levels between 30
Fig. 15.15. Light distribution in a large round- and 50% full sun; and (iii) a poorly exposed
crowned ‘Delicious’ apple tree (from Looney, 1968). zone with light levels lower than 30% full
sun. At 14 days after full bloom (DAFB) most very small portion of the canopy with poor
of the lower interior portion of the central- exposure (Fig. 15.16). At 44 DAFB, the area of
leader/M.7 trees, which had three tiers of poorly illuminated and marginally exposed
permanent branches, already had poor light canopy areas had increased in both tree
exposure (< 30% full sun), while the smaller types, but a higher proportion of the inter-
central-leader/M.9/MM.111 trees, which had stem canopy was well exposed. At 74 DAFB,
two tiers of permanent branches, only had a the relative areas of poorly and marginally
(a) 3.5 CL/M.7

< 30% Full sun
3.0 30–50% Full sun
Tree height (m)
2.5 >50% Full sun
2.0
1.5
1.0
0.5
0
14 44 74 110
(b) 2.5 CL/M.9/MM.111
< 30% Full sun
2.0 30–50% Full sun
Tree height (m)
> 50% Full sun

1.5
1.0
0.5
0
14 44 74 110
(c) 2.5
Slender spindle/M.9
2.0 < 30% Full sun
Tree height (m)
30–50% Full sun

1.5 > 50% Full sun
1.0
0.5
0
14 44 74 110
(d) 2.5
2.0
Tree height (m)
1.5
1.0 Y-trellis/M.26
< 30% Full sun
0.5 30–50% Full sun
> 50% Full sun
0
14 44 74 110
Fig. 15.16. Light distribution pattern at four times during the growing season for 11-year-old ‘Empire’ trees
trained to: (a) the central leader (CL) system on M.7; (b) the mini-CL system on M.9/MM.111 interstem; (c) the
slender-spindle system on M.9; or (d) the Geneva Y-trellis system on M.26 (from Robinson et al., 1991b).
390 T.L. Robinson
illuminated areas were smaller in the inter- much the same and only the top of the tree
stem tree than in the M.7 trees; however, was well exposed. With summer pruning,
both tree types had large areas of poorly only upright shoots were removed, but in
illuminated canopy areas. At 100 DAFB, the the compact tree of the slender spindle the
trees were summer-pruned by removing spur- and bourse-shoot leaves also account
upright shoots in both the first and the for much of the shading. With slender-
second tiers. Summer pruning improved spindle trees, the branches are usually much
light distribution in the middle section of the closer together, with only small gaps in the
canopy for both tree types. The area of canopy, than with taller trees. This results in
poorly exposed canopy in the interstem tree a high density of foliage in the tree canopy.
was reduced to a small area near the trunk With moderate to vigorous growth, these
on the bottom tier of branches. With the gaps can be closed very quickly in the season
larger M.7 trees, the area of low light level leading to poor fruit colour and quality if the
was larger and the area of marginally trees are not summer-pruned (Corelli and
exposed canopy was not reduced as much by Sansavini, 1989). Our experience with culti-
summer pruning. There was very little fruit vars like ‘Empire’ indicates that summer
in the lower interior part of the M.7 trees, pruning of slender-spindle trees is essential
while the interstem trees had fruit in all for good fruit colour. Sansavini et al. (1981)
parts of the canopy. Although summer prun- found that, under vigorous growth condi-
ing improved the light distribution of the tions, light levels in the lower part of the
M.7 canopy, it was too late in the season to canopy of a slender-spindle multi-row
have much of an effect on fruit-bud differen- system were lower than in a medium-
tiation. The improved light distribution of density palmette hedgerow. In one of our
the small central-leader tree resulted from orchard trials of five different systems, we
the smaller height and depth of the canopy found that the fruit quality and economic
than in the larger central-leader trees. Barritt returns in the eighth year were best for the Y
et al. (1991) have also reported that there is a trellis and were poorest with the triple-row
rapid decline in light exposure to less than slender-spindle system, despite it having the
20% full sun by early June for the interior highest yield. The challenge with slender-
parts of the traditional central-leader spindle trees is to combine the correct
canopy. The rapid seasonal decline in light vigour-control techniques and pruning with
exposure of the interior of central-leader spacing. Incorrect choice of tree spacing or
trees has led to modifications of the central- excessive vigour can result in excessive
leader tree form, such as the palmette leader shade, due to the limited area allotted to
(Lakso et al., 1989a), and to summer pruning each tree.
to improve the light distribution and fruit Planar canopies have been developed to
colour of most red cultivars. overcome the problems of light penetration
In the study described above (Robinson et into thick canopies. Since the foliage and
al., 1991b), the slender-spindle/M.9 trees had limbs are restricted to a single plane, these
very good light exposure to most of the tree forms usually have a dense canopy,
canopy at 14 DAFB (Fig. 15.16). All parts of which is essentially non-transmitting. The
the canopy had > 30% full sun and only a rule of thumb of 1 m of light penetration
small portion in the bottom was below does not hold in this case. With the horizon-
50% full sun. However, by 44 DAFB, light- tal planar canopies, such as the T or Ebro
exposure levels in all parts of the canopy trellis, there is a drastic reduction in light
except the top of the tree had dropped con- levels from the top to the bottom side of the
siderably, indicating that the gaps between canopy. Ferree et al. (1989) reported that,
limbs had been closed by new shoot growth. with the Lincoln canopy (T trellis), severe
By 74 DAFB, there was a large poorly illumi- dormant pruning was required for good
nated area of the canopy, which was light transmission through the canopy, with
markedly reduced by summer pruning, but more typical moderate pruning having very
the marginally illuminated area remained low transmission values. With the Ebro trellis,
Tustin et al. (1989) and Warrington et al. mette hedgerow had better light distribution
(1996) found low light transmission through into the canopy than did slender-spindle,
the top layer of the trellis, resulting in interstem or pyramid-hedgerow trees. The
excessive shade of the lower layers. These trellis also had greater crop density and
horizontal canopy systems, which were greater efficiency than the other systems. If
developed for mechanization of the harvest, canopies are kept thin, the vertical-trellis
suffer from the horticultural problems of canopies generally produce excellent fruit
excessive upright shoot growth from the top quality. However, with wide palmette-trellis
side of the trellis. This increases the level of hedgerows, Ferree et al. (1989) found light
shading and, in the case of the Ebro trellis, transmission to be similar to that of the
can result in complete closure of the space slender spindle.
between tiers. Our experience in New York In general, planting systems that result in
with red cultivars, such as ‘Empire’, shows interior canopy shading when the trees are
that fruit colour with horizontal trellises is mature, such as the triple-row or overly vig-
poor and canopy development has been orous close plantings, will have lower fruit
slow due to the flat limb orientation. quality than those that maintain good light
Inclined V- or A-shaped canopies were distribution throughout the tree canopy,
also developed for mechanical harvest, but such as the vertical axis, palmette trellis and
they do not have the vigorous-shoot prob- Y trellis. Good light distribution can be
lems of the horizontal canopies. In addition, achieved in older trees if the top of the tree is
light exposure to the bottom side of the kept narrower than the bottom of the tree
canopy depends both on light transmission and if there is a good balance between vege-
through the canopy and light energy coming tative growth and cropping. For conic
through the open gaps at the top of the systems, such as vertical axis, slender spindle
canopy arms. In our study described above, or the traditional central leader, maintaining
a dwarf version of a Y-shaped hedgerow had a conic shape as the trees age is critical to
very good light exposure at 14 DAFB, with maintaining good light exposure, fruiting
no part of the canopy receiving less than 30% and fruit quality in the bottom of the tree. In
of full sunlight (Fig. 15.16). By 44 DAFB, our experience, the best way to maintain
there was a strong gradient of light through good light distribution within the canopy as
the canopy, with the interior of the Y receiv- the tree ages is to remove whole limbs in the
ing greater than 50% full sun, while the top of the tree once they grow too long,
underside of the Y received less than 30%. By rather than shortening back permanent
74 DAFB, only the top interior portion of the scaffold branches in the tops of trees.
Y was well illuminated. Summer pruning, Traditionally with the central-leader system,
which consisted of removal of unwanted permanent tiers of upper branches were
watersprouts in the interior of the Y, developed. A successful approach to manag-
increased the exposure at the centre of the Y. ing the tops of trees has been annually to
However, the underside of the trellis remove one or two large upper branches
remained below 30% full sun. This was due completely. When this style of pruning is
to closing of the canopy of adjacent rows, repeated annually, the top of the tree can be
which allowed little light between the rows. composed completely of young fruitful
Our experience indicates that, when the branches. The younger branches do not
canopies of adjacent Y-trellis rows touch, cause as much shade as larger, older
fruit colour and quality on the underside of branches and are naturally shorter than the
the trellis decline. A minimum of 1.5 m of bottom branches, thus maintaining the conic
open space between the Y arms of adjacent shape of the tree. When this strategy, which
rows is required for good fruit colour. is termed limb-renewal pruning, is
Thin vertical canopies receive light expo- employed with high-density systems, such
sure from both sides of the canopy and so as vertical axis, slender spindle or Y trellis,
should have good light distribution within good light distribution can be maintained
the canopy. Ferree (1980) reported that a pal- over the life of the tree.
392 T.L. Robinson
15.8.5 Balance between vegetative growth that are larger than the 3 : 1 rule should be
and cropping removed early on to preserve a hierarchy of
branch and leader diameter.
The successful management of apple trees in The most important method of inducing
any high-density system depends on main- cropping and reducing induced juvenility is
taining a balance between vegetative growth tying down of the scaffold branches to
and fruiting. If vigour is too low, excessive induce cropping. In some systems, this is
fruiting results, fruit size declines, biennial limited to the first tier of branches (vertical
bearing increases and trees fail to fill their axis, slender pyramid and HYTEC), while, in
allotted space soon enough to make the other systems, upper branches are also tied
orchard profitable. If vegetative vigour is down (SolAxe). In most climates, if pruning
excessive, then flowering and fruiting are of branches is minimized, often crop load
reduced and containment of the tree to the will bend branches down and a natural
allotted space becomes problematic. The balance between vigour and cropping will be
successful balance of vegetative vigour and established without additional limb position-
fruiting results in ‘calm’ trees that produce ing. In vigorous and/or warmer climates
heavy annual crops and require only a light where winter chilling is insufficient, limbs
annual pruning. Pruning and tree-training often become too large before they set
strategies are the primary management sufficient crop loads to bend the branches
methods, along with fertilization strategies, down. In these climates, the tying down of
that are used to achieve a balance between all vigorous limbs must be done annually for
vegetative growth and cropping throughout the first 3–5 years until the tree settles down
the orchard’s life. Pruning, training and and begins to crop heavily (Cook and
fertilization strategies must be compatible Strydom, 2000). However, in most traditional
with the tree spacing, cultivar and rootstock. apple-growing areas, growers often invest
Often growers use pruning techniques that are too much money in limb positioning, which
not compatible with high-density systems. should be limited to only what is essential in
For example, pruning strategies that use the first 2 years. Thereafter, the precocity of
heading cuts can stimulate excessive tree the rootstock induces heavy cropping and a
growth and vigour, reduce fruiting and natural balance is established.
ultimately delay profitability. Management of cropping during the first
During the development years of an 4 years to avoid biennial bearing is critical to
orchard’s life the balance between vegetative maintaining a proper balance between vege-
growth and cropping of most modern high- tative growth and cropping as the trees
density systems is based on the principle of begin to bear. With precocious dwarfing
minimal pruning during the first 4 years. No rootstocks, young apple trees can often over-
heading cuts should be done to the leader set in the second or third year, resulting in
except at planting if the tree planting density biennial bearing as early as the fourth year.
requires significant tree growth to fill the This then results in increased vigour in the
allotted space. This heading cut serves to fourth year, just when the trees have filled
balance the top of the tree with the root sys- their allotted space and when reduced
tem to ensure good growth in the first year. vigour is needed. Cultivars differ in their
Thereafter, the maximum growth is achieved biennial bearing tendency and this must be
with no pruning. For the first 4 years, incorporated into the crop loads allowed on
pruning should be limited to the removal of young trees. In general, we recommend crop
unsuitable branches, such as those lateral loads of five to 15 apples per tree in the
branches that are as large as or larger than second year, 30–60 apples per tree in the
the leader. Cook and Strydom (2000) have third year and 100–120 apples per tree in
suggested a 3 : 1 pruning rule where the the fourth year. Within each year, the low
leader should be three times the diameter of end of the range should be used for low-
any of the lateral branches in the upper part vigour trees and the high end of the range
of the tree. They have suggested that limbs for high-vigour trees.
As the orchard reaches maturity, contain- 15.9 Orchard Management Practices for
ment pruning of the canopy is critical to Successful High-density Orchards
maintaining trees within the allotted space.
Pruning strategies based on shortening or There are many additional factors that must
stubbing back permanent branches that out- be considered to make the new high-density
grow their allotted space are not generally as orchard truly profitable. In one of our studies,
successful as limb-renewal pruning strategies. we obtained yield data from ten grower
This is partially because the most productive orchards of the same high-density vertical-
fruiting wood is cut off when a branch is axis system. There was considerable variation
shortened. In addition, stubbing cuts stimu- in early yields. Those orchards with low
late localized vigour on the affected branches. yields could almost always be traced to poor
In our studies on how to manage the canopies tree growth in the first 3 years, which
of both low- and high-density systems, treat- delayed the time when the trees achieved
ments where branches were shortened to canopy closure. If trees do not grow well in
maintain the conic shape of the tree resulted the first few years, excessive flowering and
in unacceptable yield reductions and exces- fruiting occur, which stunts the tree. Such
sive vigour compared with an unpruned con- orchards often fail to fill the allotted canopy
trol. A more successful approach has been to space and continue to have moderately low
annually remove one or two large upper yields for many years. Management variables
branches completely and develop younger that have a large impact on the growth and
replacement branches. To ensure the develop- performance of young trees over the first 5
ment of a replacement branch, the large years include the following.
branch should be removed with an angled or
bevelled cut so that a small stub of the lower
portion of the branch remains. From this stub 15.9.1 Soil and site selection
a flat weak replacement branch often grows.
If these are left unheaded, they will naturally Site and soil factors initially determine
bend down with crop. They are naturally whether an orchard should be planted at all
shorter than the bottom branches, thus main- (see Chapter 11). There are many orchards in
taining the conic shape of the tree without existence today that would not have been
stubbing cuts. This type of pruning does not planted had growers carefully considered the
stimulate vigorous regrowth. Using the 3 : 1 real costs of planting in that site. Site and soil
pruning rule of Cook and Strydom (2000), characteristics that result in crop or tree loss
where limbs that are larger than one-third the rapidly reduce the profitability of an orchard
size of the leader are removed, can result in a block. This is especially true with high-den-
tree with primarily small fruiting branches in sity orchards. If the site selected has the
the top of the tree. Our recommendation is to potential for spring frosts, winter freezes,
begin removing one or two whole limbs in fruit russet, tree loss or hail, resulting in crop
the top of the tree once the tree is mature loss during these early years, then the new
(year 6–7). This allows moderate pruning high-density block will not be profitable.
each year and is a method to contain tree Unsuitable sites cannot usually be modified
size. It also maintains good light distribution economically to prevent finish and fruit-qual-
in the canopy without inducing excessive ity problems. In almost all cases, if growers
vigour. On trees with overgrown tops that planted only their best sites with new high-
need to be restructured, moderate renewal density orchards and left the undesirable
pruning (one or two large upper branches sites unplanted, their profitability would be
annually) for a 4–5-year period can eliminate improved. Often significant amounts of
all of the large branches in the top of the tree. money are wasted on poor sites. Soil charac-
This style of pruning can be applied to teristics that result in excess water or lack of
almost all systems. Even those systems based water are also important, but can sometimes
on permanent branches can benefit from be modified through choice of rootstock or
renewal pruning as they age. mechanical methods, such as drainage, irri-
394 T.L. Robinson
gation or ridging. The fruit grower needs to 15.9.3 Time of planting

know whether the cost of modification will
bring economic returns. ‘Soil strength’ or fer- Late planting has been shown to reduce tree
tility can sometimes be important, especially growth in the first year compared with early
with vigorous and poor-colouring cultivars. spring or autumn planting (Robinson and
Excess fertility often results in soft and Stiles, 1991). To maximize tree growth in the
poorly coloured, unmarketable fruit. This first year, growers should plant as early in
fruit is not suitable for extended storage or the spring as possible and, in climates where
export and will result in lower profitability, this is possible, trees should be planted in
since alternative markets are less profitable. the autumn.
Alternatively, if soil fertility is low and trees
grow poorly for the first few years, then prof- 15.9.4 Weed control
itability will be significantly reduced.
Weed competition can drastically reduce tree
growth during the first few years and can
15.9.2 Tree quality cause a failure of the orchard to fill its allot-
ted space, which always results in dimin-
Several studies have shown that larger caliper ished yield and profitability. Merwin and
trees grow more and produce larger yields in Ray (1997) have shown that good weed con-
the first 4–5 years than do smaller caliper trees trol during the first 3–4 months of a growing
(Van Oosten, 1978; Ferree and Rhodus, 1987; season is the most critical time period of the
Robinson and Stiles, 1991; Sanders, 1993; season. In later summer months, if weed
Wertheim et al., 1995). Large caliper trees that control is poorer it is not detrimental to the
also have lateral branches or feathers produce trees. Thus we recommend that growers
more than unbranched whips, especially in provide excellent weed control for the first
the second and third year (see also Chapter 6). 4 months of the season for the development
For very high-density systems that depend on years of the orchard (see also Chapter 13).
significant second- and third-year yield, feath-
ered trees are vital to the economic success of
15.9.5 Irrigation
the system. If growers use small-caliper whips
that do not produce fruit until year 4 or 5,
In many years, dry weather following plant-
often the carrying costs from the extremely
ing results in water stress of newly planted
high investment of high-density orchards trees, which can limit tree growth. Often trees
overwhelm the potential returns and negate undergo stress despite moderate soil-water
the benefit of the high tree density for profit- levels. This is due to the damaged root system
ability. We recommend that the caliper of trees of a transplanted tree, which cannot ade-
used in high-density plantings be a minimum quately support the top without frequent irri-
of 15 mm and that they have five to ten well- gation. This is especially true with
positioned feathers, with a maximum length large-calliper feathered trees. Feathered trees
of 30 cm and starting at a minimum height of produce significant leaf area shortly after
80 cm on the tree. An exception to this recom- planting, which develops a high water
mendation is with the super-spindle or V demand before the root system can regrow
super-spindle system, where the cost of such sufficiently to support the trees. Frequent
high-quality trees makes these systems unprof- trickle irrigation can help these trees produce
itable. With these systems, much cheaper trees good growth in the first year. In humid areas,
are needed, and either sleeping-eye budded growers are unaccustomed to installing irriga-
rootstocks or medium caliper trees with many tion or delay its installation until midsummer.
short shoots are preferred. With these systems, We recommend that growers install trickle
the trees are planted so close together irrigation soon after planting with high-den-
(50–60 cm) in the orchard that very little addi- sity orchards that use feathered trees, in order
tional lateral extension growth is needed after to prevent water stress and maximize tree
planting to fill the space. growth.
15.9.6 Fertilization 15.9.7 Soil fumigation
Frequent low doses of nitrogen fertilizer, In many cases, tree growth of new orchards
either through the trickle system or on the that are planted on old orchard land can be
ground with irrigation, will generally improved significantly with soil fumigation.
improve tree growth during the first few However, the apple replant problem is vari-
years to speed development of the canopy. able, with some sites showing no benefit
Excessive fertilizer applications near the from fumigation and others showing signifi-
tree trunk should be avoided, since they cant benefits (Merwin et al., 2001). Ideally,
can stunt tree growth in the first year due growers should conduct a bioassay before
to salt toxicity. Excessive fertilization, espe- replanting an old orchard site in order to
cially nitrogen, can cause too much growth, assess the severity of the replant problem
which results in greater pruning costs, and determine the value of soil fumigation.
delayed flowering, reduced yields and poor Even with fumigation, almost all old orchard
fruit quality. Either too little tree growth or sites produce less tree growth than virgin
too much tree growth will result in failure sites. Thus, tree-planting density should be
to achieve expected yields and in decreased increased on old orchard sites compared with
fruit quality. Nitrogen management of virgin sites by 20–30%.
high-density orchards must take into con- If growers properly combine the manage-
sideration tree-planting density. For low ment variables listed above, they should
tree densities, trees must be grown vigor- obtain 50 cm of leader shoot growth in the
ously for several years to fill the allotted first year, 75–100 cm of leader shoot growth
space with canopy, and relatively high in the second year and 50 cm of leader shoot
nitrogen fertilization is desirable for several growth in the fourth year. If this is combined
years after planting. However, as tree den- with minimal pruning and a precocious root-
sity is increased, fewer years are required stock, significant production should be
for the tree canopy to fill the allotted space. obtained in the third and fourth years, which
In the case of moderately high-density sys- will limit vegetative growth in future years,
tems, such as vertical axis, slender pyramid resulting in a ‘calm’ tree.
or Y trellis, high nitrogen fertilization
should only be used during the first 2
years, while the canopy is still expanding. 15.10 Economic Comparisons of
With the higher tree densities used in the Orchard Planting Systems
slender spindle or V slender spindle, there
is little need for additional lateral tree The economic performance of different
growth after planting when highly orchard systems is the final arbiter of the
branched trees are used, and any vigorous value of a system. Economic measures that
tree growth is counter-productive. In the are important to fruit growers are the costs
extreme densities of the super-spindle sys- of establishment, the cost of mature orchard
tem, no new lateral canopy extension is maintenance, the time required to pay back
needed and nitrogen fertilization in the first the investment, the annual cash flows, the
few years is avoided in order to enhance discounted cash flows, the net present value
cropping and settle the trees down. In of the investment, the annuity of the net
general, growers should only grow trees present value and the expected life of the
vigorously until the canopy fills the allotted orchard. An additional factor, which is diffi-
space. Thereafter, low nitrogen fertilization cult to quantify, is the level of financial risk
is desirable to keep the trees calm with a associated with each system.
balance between fruiting and cropping. Most economic studies have shown
Many mature high-density orchards receive greater investment costs and annual labour
excessive nitrogen fertilizer rates, which costs with increasing tree density. However,
cause severe canopy management problems due to higher early yield and higher cumula-
(see also Chapter 12). tive yield, profitability is generally increased
396 T.L. Robinson
with increased tree density (Goedegebure, cost of trees and support was only 20% of
1978, 1980, 1986, 1989). However, due to the the total investment required. With the
law of diminishing returns with additional higher-density systems, the maximum
trees per hectare, extremely high tree densities investment required was higher than with
have been shown to be less profitable than the central-leader system, with the V-slen-
more moderate densities. In addition, risk der-spindle system requiring an investment
increases with increasing investment, making of US$41,400 ha1. As tree density was
the very high-density systems higher-risk. increased, the percentage of the total invest-
We have evaluated the economic per- ment accounted for by tree and trellis costs
formance of seven orchard planting systems also increased. Thus, efforts to reduce tree
(central leader/M.111, mini-central leader/ cost and/or support-system cost would
M.9/MM.111, palmette trellis/M.9, Y trellis/ have little impact on the total investment
M.26, vertical axis/M.9, slender spindle/ required to plant the low-density central-
M.9, V slender spindle/M.9), using yield and leader system, but would have a large
cost data from our research plots in New impact on the higher-density systems, espe-
York State (White and DeMarree, 1992; cially the V slender spindle. The cumulative
DeMarree, 1995a,b; Robinson et al., 1996b). cash-flow curve did not break even until the
The systems represent a range of tree densi- tenth to the 13th year, depending on the sys-
ties, from 500 to 2245 trees ha1, and a range tem. The vertical-axis, Y-trellis and palmette-
of rootstock vigour, from MM.111 to M.9. trellis systems had the quickest break-even
The systems varied in costs from US$7370 time and the central-leader system the slow-
for the central-leader system to US$28,460 est. Thus, although the low-density central-
for the V-slender-spindle system (Table leader system had the lowest initial
15.11). The large differences in establishment investment requirement, it had a longer
costs were largely related to tree density and break-even period than did all of the higher-
the requirement of support for the dwarf density systems. After 20 years, the Y-trellis,
rootstocks used in the high-density systems. vertical-axis and slender-spindle systems
Early yield and the maximum yields had the greatest cumulative cash flows, with
achieved by each of the seven systems also more than double the cumulative cash flow
varied widely. The V-slender-spindle sys- of the central-leader system.
tem reached its maximum yield of 42 t ha1 From an investment-analysis perspec-
by year 6, while the central-leader system tive, systems can be compared by calculat-
reached its maximum yield of 35 t ha1 by ing the internal rate of return on the
year 10. The speed at which each system investment over the 20-year life of the
reached its maximum yield was related to orchard or by discounting the annual net
the tree density and to the precocity of the cash flows back to the present value, using
rootstock. a fixed discount rate (net present value
The systems varied widely in how nega- (NPV)). We have assumed a real interest
tively the cumulative cash-flow curve rate (or discount rate) of 6%. All of the sys-
dipped and when the cumulative cash flow tems had a positive internal rate of return
curve became positive (Table 15.11). (IRR), which ranged from a low of 8.2% for
Although the low-density central-leader the central-leader system to a high of 11.8%
system required the least investment in tree for both the vertical-axis and the Y-trellis
and establishment costs, the system systems (Table 15.11). The slender-spindle
required a total investment of US$23,100 system and the palmette trellis were
ha1 by the end of year 5 before the cumula- intermediate at 10%. In general, the higher-
tive cash-flow curve turned upwards. Many density systems had the highest IRR, with
growers have not fully realized that the ini- the exception of the highest-density, V-
tial investment in trees and support is only slender-spindle system, which had a similar
a fraction of the total cost of establishing an IRR to that of the central-leader system.
orchard and bringing it into production. In Examined from the perspective of NPV of
the case of the central-leader system, the the accumulated cash flow over 20 years,
Apples - Chap 15
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3:31 pm
Table 15.11. Costs and profitability of seven orchard systems in New York State (from Robinson et al., 1996b).
Year Net present

of most Internal value of
Total Maximum negative rate of accumulated Net Annual
Page 397
Tree 1st-year negative cumulative return profit present- equivalent
Apple-orchard Planting Systems

density costs cash flow cash flow over over value cash flow
(trees (US$ (US$ (US$ 20 years 20 years break-even (US$
System/rootstock ha1) ha1)a ha1) ha1) (%) (US$ ha1) year ha1)
Central leader/M.7, 500 7,369 23,117 5 8.2 5,024 17 437

MM.106 or MM.111
Mini-central leader/ 680 9,351 24,227 5 10.4 11,476 15 1001
M.26 or interstems
Vertical axis/M.26, 1280 17,908 29,670 3 11.8 19,732 13 1720
interstems
Y trellis/M.26, interstems 1280 18,150 29,937 4 11.8 19,876 13 1754
Slender spindle/M.9 1500 20,164 34,130 4 10.9 17,943 14 1564
Palmette trellis/M.9 1500 17,016 29,181 4 10.7 14,686 14 1280
V slender spindle/M.9 2245 28,465 41,406 3 8.8 10,594 16 924
a Includes land cost of $2000 ha1 and pre-plant land preparation.
397
398 T.L. Robinson
our analysis showed that all systems were tral leader, mini-central leader, palmette
profitable. The NPV of accumulated profit trellis and V slender spindle) (Table 15.12).
after a 20-year life of each orchard system All of the high-density systems are more
ranged from a high of US$19,876 for the Y- sensitive to price than the low-density cen-
trellis system to a low of US$5024 for the tral-leader system. This means that under
central-leader system. With this method of low prices they drop the most, but also
analysis, the central-leader system was under high prices they benefit the most. A
clearly the least profitable system. The high- good example is the situation when a new
density systems divided into two groups. cultivar brings high prices for the first few
The most profitable group was the Y-trellis, years. At US$0.52 kg1 during the first 10
vertical-axis and slender-spindle systems, years, the best high-density systems (Y trel-
while the high-density central-leader, pal- lis, vertical axis and slender spindle) would
mette-trellis and V-slender-spindle systems produce triple the profits of the low-density
were intermediate in profitability. central-leader system. Under low-price sce-
There were large differences in the year narios, the low-density system would
when the different systems broke even (Table become unprofitable before the best high-
15.11). The earliest break-even year was year density systems.
13 for the vertical-axis and Y-trellis systems. If the yields of each system were reduced
The latest to break even was the central by 5 t ha1 each year, then only the central-
leader (year 17). The break-even year indi- leader system would no longer be profitable
cates the minimum lifetime of the different (Table 15.12). However, if yields were
systems. Thus, the vertical axis and Y trellis reduced by 10 t ha1 each year, then only
would need to last at least 13 years to recoup the three best systems would be profitable
the investment and earn 6% on the invest- (Y trellis, vertical axis and slender spindle).
ment, while the central leader would need to If yields were reduced by 15 t ha1, then all
last at least 17 years. Systems with a shorter of the systems would be unprofitable.
break-even period allow the grower flex- Under this scenario the lowest-density sys-
ibility of replanting to a different cultivar tem and the highest-density system would
should market conditions change. lose the most money. In general, the prof-
Another interesting comparison is the itability of the high-density systems is more
annual contribution to orchard cash flow, in sensitive to yield than the low-density cen-
discounted dollars (annual equivalent cash tral-leader system.
flow (AECF)) each system would make If land prices were increased from the rel-
(Table 15.11). This measure allows the calcu- atively low price in western New York of
lation of the area required to produce a US$2000 ha1 to US$12,000 ha1, the profit-
US$100,000 AECF. According to our analy- ability of all systems would be reduced, but
sis, 228 ha of the central-leader system only the low-density central-leader system
would produce a similar AECF to only would become unprofitable (Table 15.12).
57–64 ha of the three best systems (Y trellis, In a Dutch study of commercial slender-
vertical axis and slender spindle). spindle orchards at different densities,
Regardless of which high-density system is investment cost and labour hours required
chosen, a grower would need less than half for planting, pruning and tree training were
the area of high-density orchards com- increased with increasing tree density
pared with the central-leader system to pro- between 2000 and 4000 trees ha1
duce a US$100,000 AECF. (Goedegebure, 1991, 1993). These increased
These results are based on obtaining the costs were offset by increased cumulative
yields and prices we have assumed. It is yield with increasing tree density between
important to determine how the results 2000 and 4000 trees ha1. The economic
would change if lower prices or yields were profitability as measured by the NPV of the
obtained. If fruit prices were reduced from discounted cash flow over the lifetime of the
US$0.34 kg1 to US$0.26 kg1, four of the orchards was improved with increasing tree
systems would no longer be profitable (cen- density (Table 15.13).
Apples - Chap 15
Table 15.12. Effect of farm-gate fruit price, yield and land price on profitability of seven orchard systems in New York State (from Robinson et al., 1996b).
NPV (US$) NPV (US$) NPV (US$) NPV (US$) NPV (US$)
with with high with with with
21/3/03
standard NPV (US$) fruit price reduced reduced reduced NPV (US$)
fruit price, with low for first yields yields yields with high
yield and fruit price 10 years (reduced (reduced (reduced land cost
System land costa (US$0.26 kg1) (US$0.52 kg1) 5 t ha1) 10 t ha1) 15 t ha1) (US$12,000 ha1)
3:31 pm
Central leader/M.7, 5,024 7295 12,404 927 7475 14,676 2,301
MM.106 or MM.111
Mini-central leader/ 11,476 2340 22,066 5,298 1530 8,681 4,161
M.26 or interstems
Page 399
Vertical axis/M.26, interstems 19,732 1994 43,123 11,718 3492 5,429 12,420

Y-trellis/M.26, interstems 19,876 1334 40,981 11,822 3810 4,809 12,566
Slender spindle/M.9 17,943 72 39,147 9,894 1147 7,715 10,631
Palmette trellis/M.9 14,686 1557 34,949 5,891 2870 11,886 7,374
V slender spindle/M.9 10,594 8429 36,429 1,737 7351 17,122 3,282
a Standard farm-gate fruit price = US$0.32 kg1; standard land cost = US$2000 ha1; standard mature yield for central leader = 35 t ha1, for high-density
central leader = 38 t ha1, for vertical axis = 42 t ha1, for Y trellis = 45 t ha1, for slender spindle = 42 t ha1, for palmette trellis = 38 t ha1 and for V slender
spindle = 42 t ha1.
Table 15.13. Labour requirements and economic performance of Dutch slender-spindle orchards at three densities (from Goedegebure, 1993).
Cost of Pruning
Tree trees and Labour Pruning Training Pruning Pruning Pruning Pruning Pruning h ha1
density support h ha1 h ha1 h ha1 h ha1 h ha1 h ha1 h ha1 h ha1 (year 7 NPV
ha1 (Dfl ha1) planting (year 1) (year 2) (year 2) (year 3) (year 4) (year 5) (year 6) and up) (Dfl ha1)
2000 16,680 189 9 45 9 23 47 62 66 91 169,000

3000 22,440 232 11 80 17 31 56 76 77 102 196,000
399
4000 27,920 275 12 116 25 39 66 90 89 114 223,000
400 T.L. Robinson
Weber (2000, 2001) reported an economic 15.11 Conclusions

study of the slender-spindle and super-
spindle systems from commercial orchards The key objective in planning a new orchard
as grown in the Bodensee area of Germany with any modern planting system should be
(Table 15.14). The slender-spindle system to maximize yield in the early years and still
had a density of 3000 trees ha1 and the effectively produce large yields of high-
super-spindle system 6429 trees ha1. The quality fruit after the trees are mature. The
maximum yield per tree was estimated to best way to obtain high early yields is to plant
be 13.5 kg per tree (40.5 t ha1) in year 5 high tree densities, irrespective of the tree
with the slender spindle and 7.7 kg per tree shape or training system. Successful tree den-
(49.5 t ha1) in year 4 with the super spin- sities under current economic conditions have
dle. The super-spindle system used less ranged from 1000 to 7000 trees ha1. The opti-
expensive trees (0.63 Euro less per tree) than mum tree density is primarily an economic
the slender-spindle system. Nevertheless, at issue. In most apple-producing areas, the opti-
the end of the first year, the cash-flow value mum economic density appears to be between
of the super-spindle system was 6614 Euros 1000 and 3000 trees ha1. In some areas with
more negative than for the slender-spindle very high land prices, very low tree prices,
system. The higher yield of the super-spin- very high fruit prices and/or government sub-
dle system resulted in a break-even cash sidies for planting, the optimum economic tree
flow 1 year later. At the end of the 10th year, density appears to be from 3000 to 6000 trees
the super-spindle orchard had a 2306 Euro ha1. In all areas of the world, the traditional
ha1 advantage in NPV over the slender- low-density central-leader system is not as
spindle orchard. The super-spindle system profitable as the higher-density systems. In
also had a higher picking output of 30 kg addition, the low-density central-leader
h1 and reduced tree-management labour system generally becomes unprofitable before
cost, including tree training and hand-thin- the high-density systems under poor price or
ning, of 89 h ha1. yield scenarios. The super-high-tree-density
systems have substantially greater risk than
Another Dutch study compared trees at
moderately high-density systems, due to the
densities from 3000 to 20,000 trees ha1.
very high investment costs. Often they are
Although the best production was achieved
only marginally more profitable than more
between 10,000 and 20,000 trees ha1 (Fig.
moderate high-density systems, such as verti-
15.13), the best average annual cash flow of
cal axis, slender spindle, slender pyramid,
‘Jonagold’ or ‘Elstar’ on M.9 rootstock was
HYTEC or Y trellis. Regardless of the system,
obtained with a tree density of 6000 trees
growers must understand the high investment
ha1, followed by a density of 3000 trees
cost of planting any high-density system and
ha1 (Table 15.15). Profitability was lower
should limit planting cost and limit the
with a density of 10,000 and was negative
amount of replanting to only 4–5% of their
with a density of 20,000 (Balkhoven-Baart,
total area in order to maintain a positive cash-
2000; Groot, 1997). Fruit size and quality flow position for their business.
were also reduced at the highest tree densi- Among training systems, the V systems
ties (Wagenmakers et al., 1994). An examina- have the highest yields and allow a good bal-
tion of the effect of fruit price showed that, ance between vegetative growth and crop-
with prices below 75 cents kg1, the lower ping. They generally have higher light
density of 3000 trees ha1 was the most interception than do conic-shaped systems
profitable, while, at very high fruit prices of or flat planar systems. They also provide bet-
US$1.05 kg1, the higher density of 10,000 ter light exposure on both sides of fruit than
trees ha1 had the highest profitability. If do horizontal branches. V systems also lend
tree price was very high, it favoured the themselves to very close in-row spacings and
lower densities, while very low tree prices highly rectangular planting designs, which
favoured the higher densities. allow for the use of existing equipment
Apples - Chap 15
21/3/03
3:31 pm
Page 401
Table 15.14. Economic comparison of slender-spindle and super-spindle orchards in the Bodensee area of Germany (from Weber, 2000).
Total Cash flow

Tree Tree Price establishment Years Mature Picking Training Hand- at end of
density spacing per tree cost to full yield output h ha1 thinning year 1
Training system ha1 (m) (Euros) (Euros ha1) yield (t ha1) (kg h1) (years 1–5) h ha1 (Euros ha1)
Slender spindle 3000 1.0 3.0 3.17 14,541 5 40.5 130 186 50 18,812
Super-spindle 6429 0.5 2.8 2.54 20,744 4 49.5 160 111 36 25,426
401
402 T.L. Robinson
Table 15.15. Economic performance of ‘Jonagold’/M.9 and

‘Elstar’/M.9 slender-spindle trees at various densities in The
Netherlands (from Balkhoven-Baart et al., 2000).
Tree density ha1 Annuity net present value (Dfl ha1)
3,000 7,464
6,000 10,853
10,000 3,483
20,000 5,489
in high-density orchards. Despite these the orchard is mature. This can be accom-
advantages, the higher initial costs of the plished by ensuring that row spacing is not
V systems gives them similar profitability to too wide and that tree height is at least 80%
conic systems, such as vertical axis, HYTEC, of row spacing. High fruit quality when the
slender pyramid or slender spindle, when orchard is mature requires good light distrib-
planted at the same density. The greater ution throughout the tree canopy. This can
simplicity of conic systems has meant that best be accomplished by maintaining a nar-
most growers are opting for the conic-shaped row canopy shape and through regular limb-
systems. V systems will probably be better renewal pruning.
than conic-shaped tree systems under condi- In the future, the most successful apple
tions of high sunburn and high winds or growers in the world will need to produce
where all the fruit must be picked from the apples very efficiently. This will be done by
ground, since the V systems can intercept planting new high-density orchard systems
more light with short-stature systems than with new higher-quality cultivars. It appears
pyramid-shaped systems. that most of the new orchards will be devel-
All high-density systems depend on early oped with a vertical conic-shaped tree using
yields to be profitable. Growers should focus M.9 rootstock or M.9-size fire blight-resistant
on achieving high early yields by selecting rootstocks. Tree height will be 3–4 m, which
precocious rootstocks and frost-free sites and maximizes light interception at maturity,
by obtaining excellent early tree growth. If with 3–4.5 m between rows. Successful grow-
this is combined with the proper tree training ers will rely on minimal pruning of the leader
and minimal pruning, high-density systems and the scaffolds in the early years, which
can be much more profitable than traditional will help to maximize early yield. They will
systems. The specific tree-training recipe utilize a simple support system and a tree-
used in the development of an orchard is less training recipe that is simple and easy to
important than the planting density, as long teach to unskilled labour. Lastly, they will
as minimal pruning is employed. manage the canopy of mature trees to ensure
High yield per unit land area at maturity good light distribution within the canopy in
requires high total light interception when order to maximize yields and fruit quality.
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16 Flower and Fruit Thinning and

Vegetative : Fruiting Balance
Ross E. Byers
Department of Horticulture, Virginia Polytechnic Institute and State University,
Winchester, Virginia, USA

16.2 History of Flower and Fruit Thinning 410
16.3 Commercially Important Physiological Effects of Thinning 410
16.3.1 Return bloom 410
16.3.2 Fruit size, quality, colour and pest control 411
16.3.3 Yields 413
16.3.4 Defruiting young trees for tree growth 413
16.3.5 Effect of crop reduction on growth and fruiting of bearing trees 414
16.4 Thinning Chemicals and Their Physiological Effects 414
16.4.1 Pollinicides 415
16.4.2 Gibberellin for inhibition of flower-bud formation 416
16.4.3 Hormone-type plant-growth regulators (PGR) for bloom and fruit thinning 416
16.5 Adjuvants 421
16.6 Application Considerations 422
16.6.1 Drying rate (dew/rain/humidity/rewetting) 423
16.6.2 Effect of application temperature on chemical fruit thinning 423
16.7 Influence of Photosynthetic Reserves on Fruit Set and Chemical Thinning of Apples
Following Ovule Fertilization 424
16.7.1 Effect of light and darkness on apple fruit retention and thinning 424
16.7.2 Effect of post-application temperature on chemical fruit thinning 426
16.7.3 Physiological vs. temperature basis for timing chemical applications 427
16.7.4 Temperature probabilities for thinning 428
16.7.5 Pruning and mechanical fruit thinning 428
16.8 Chemicals that Induce Flower-bud Formation Without Thinning 429
16.9 Conclusion 430
16.1 Introduction maintain tree structure. Thinning also

provides more consistent annual yields,
Fruit or flower thinning is commercially which in turn allow the efficient use of pack-
practised in order to maximize crop value aging, grading, storage equipment and
by optimizing marketable fruit sizes, yields, labour. Before the development of chemical
fruit colour, shape and quality, as well as to thinners, hand-fruit-thinning was routinely
promote return bloom and tree growth and practised. However, as labour costs increased

410 R.E. Byers
and crop values decreased, hand-thinning previous season (‘York Imperial’ and ‘Fuji’).
became less economic. Currently, For these reasons, thinning and cultural prac-
hand-thinning is used primarily to supple- tices for each individual cultivar must take
ment chemical thinning – for example, to into account the cultivar’s genetic potential
promote higher fruit quality by removal of and the climate of each production region.
small or misshapen fruit (Williams, 1979). In a strongly alternate-bearing cultivar
To a processor, apple value is frequently (‘Marigold’), Darbellay (1998) found that the
based on increasing increments of 0.6 cm in highest sustained yields and maximum crop
fruit diameter. Fruit diameter over 6.4 cm is loads of high-quality fruit were best attained
typically used as a benchmark, because larger by proper thinning for crop-load control and
fruit have a lower proportion of core and peel light pruning to ensure adequate spur and
than smaller fruit. In some years, a 10% ‘pre- shoot vigour. If trees were not pruned, exces-
mium’ price may be paid for large volumes if sive fruit thinning was required to avoid
80% of the fruit are over 6.4 cm in diameter. In biennial bearing, low yields and losses due
other years, fruit below 6.4 cm may have little to excessively large fruit. However, severe
or no value or are bought at juice price. Juice pruning alone as a method of reducing crop
price is frequently only 30% of that of larger load was not found to be a viable alternative
fruit (6.4 cm or greater) that are used for sauce to fruit thinning, due to poor yields, fruit
or slices. Fresh-marketed fruit also have a clustering and variable fruit size.
greater value if fruit diameters are within the
range of 6.4–8.9 cm. In a 5-year integrated
orchard-management study, researchers in 16.2 History of Flower and Fruit
North Carolina found that only 41% of Thinning
‘Delicious’ apples met the US fancy-grade
standards (Shaffer et al., 1983). The major col- Over 2000 years ago, Theophrastus (see
lage factor was insufficient fruit size (21%), fol- Thompson, 1957) reported that farmers recog-
lowed by insufficient colour (13.1%). nized the tendency of fruit trees to over crop
Inadequate fruit thinning soon after bloom and described the custom of partial crop
was considered responsible for excessive crop removal. In 1919, Bedford and Pickering con-
loads, which contributed to both small fruit vincingly showed that alternate bearing could
size and insufficient colour. Their data demon- be controlled by hand-thinning at the bloom
strated the major economic impact associated stage instead of fruit thinning 6–8 weeks after
with excessive crop loads (Shaffer et al., 1983). full bloom (AFB), which was the commercial
No data were presented to reflect crop losses practice at the time. In 1934, Auchter and
in subsequent year(s) due to poor return Roberts made the first conscious effort to
bloom, alternate bearing or poor tree growth. totally eliminate flowers with common spray
The inherent genetics of each cultivar are materials of the period (calcium or sodium
uniquely different for fruit size, colour, shape, polysulphide, copper sulphate, oil emulsion,
natural fruit set, return bloom and thinning zinc sulphate and tar distillates), but defolia-
responses to chemicals. Recognizing the indi- tion and fruit injury were significant prob-
vidual characteristics of a cultivar is lems with most of the chemicals tested.
extremely important for maximizing crop
value. Certain cultivars are inherently large
(such as ‘Twenty Ounce’, ‘Mutsu’ 16.3 Commercially Important
and ‘Jonagold’), while others (‘Jonathan’, Physiological Effects of Thinning
‘Winesap’ and ‘Gala’) may not achieve 6.4 cm
in diameter even when trees are moderately 16.3.1 Return bloom
cropped. Additionally, some cultivars, even
though heavily cropped in one year, may pro- The timing and severity of thinning in one
vide an excellent return bloom and crop regu- year may strongly influence cropping for
larly (‘Gala’), while other cultivars may not several subsequent years. Singh (1948)
flower the following season if trees flowered reviewed several studies and suggested that
heavily and carried a moderate crop in the fruit thinning practised 30 or more days after
Flower and Fruit Thinning 411
bloom was rarely successful for control of days or FB + 39 days. Also, larger fruit that
alternate bearing. Strongly biennial-bearing matured earlier were more susceptible to rot-
cultivars may routinely be overthinned in ting by Gloeosporilum perennans.
order to achieve return bloom, even though Large fruit from thinned trees are gener-
greater yields of adequate fruit size could be ally more susceptible to bitter pit and inter-
realized with an earlier thinning time nal breakdown and have lower calcium and
(Harley et al., 1934, 1942; Singh, 1948). higher potassium concentrations than do
Comparison studies of the effects of thinning small fruit from non-thinned trees. Further-
at bloom vs. 30 days AFB on the subsequent more, fruits from thinned trees were found
season’s return bloom, yield and fruit size to be less firm and have higher soluble-solids
were not conducted in the 1940s because concentration than those from unthinned
hand-thinning at bloom or petal fall was trees (Sharples, 1964, 1968; Volz et al., 1993).
commercially impractical. Lafer et al. (1999) determined that the
Fruit thinning, if sufficiently severe, as optimum crop load should range between
late as 60 days AFB was shown to increase 1 and 1.5 kg cm2 TSCA (6–9 fruits cm2) for
blossom-bud differentiation (Haller and several cultivars. Johnson (1992) suggested
Magness, 1933). However, some strongly that growers should be cautioned against
biennial bearing cultivars, when bloom- long-term storage of fruit from light trees or
thinned, may not have an adequate return overthinned trees and that additional cal-
bloom especially when the trees carry a full cium sprays may be needed to reduce stor-
crop to harvest (e.g. ‘York Imperial’/M.26 age disorders in some cultivars.
and ‘Golden Delicious’/M.26) (Byers et al., Volz and Ferguson (1999) found that
2000b). McArtney et al. (1996) found that bloom-thinning trees to a single fruit/spur
if ‘Royal Gala’ (an annual-bearing cultivar) greatly increased fruit size (65%) and had no
was fruit-thinned 3–4 weeks after bloom, effect on internal calcium concentration.
fruit weight was 16% less at harvest and leaf However, in contrast, thinning alternate clus-
area per tree was depressed by 17%. ters only slightly increased fruit size (21%)
and reduced calcium concentration by up to
22%. The reduced calcium concentration was
16.3.2 Fruit size, quality, colour and caused by an increase in multi-fruited clus-
pest control ters and by lower leaf areas on bearing
spurs. For cultivars that suffer from calcium
Typically, with alternate-bearing cultivars, disorders, 1-naphthyl-N-methylcarbamate
thinning in the ‘on year’ increases yields of (carbaryl) may be a more desirable thinner
the more valuable fruit sizes (Preston, 1954). since it is a more selective thinner for
In the ‘off year’, fruit in the larger size removal of smaller fruit from the cluster than
catagories (over 75 mm) frequently com- is napthaleneacetic acid (NAA), ethephon or
mand a lower price because they are too 6-benzyladenine (6-BA).
large and have more physiological disorders In ‘regular-bearing’ cultivars, total yields
and a shorter storage life. In the ‘on year’, a over a 2-year cycle may be reduced by thin-
significant portion of the fruit are in the ning, but the increased fruit size improves
smaller, low-priced categories. In large- crop value (Preston, 1954; Quinlan and
fruited cultivars, early thinning may result in Preston, 1968). Where a theoretical value is
oversized fruit at harvest and thinning may, assigned to each fruit size, the total crop value
therefore, be deliberately delayed in order to is disproportionally influenced by the larger
reduce fruit size. This delay may improve fruit (Table 16.1). In this case, crop value was
fruit firmness but have a negative impact on higher even though total fruit yields were
return bloom (Johnson, 1992, 1995). Johnson lower due to overthinning at FB + 1 week. For
(1995) reported earlier fruit maturation (as these reasons, the objectives of thinning
determined by internal ethylene, respiration should be to: (i) eliminate the smallest fruit; (ii)
rates and colour) when trees were thinned at maximize the production of the most valuable
full bloom (FB) + 5 days, but not at FB + 27 fruit sizes; and (iii) prevent biennial bearing.
Apples - Chap 16
412
11/4/03
11:01 am
Table 16.1. Percentage crop, by weight, in five size grades, and yield of ‘Sunset’/M.9 trees (2-year mean, 1964/65) (from Quinlan and Preston, 1968).
Scenario 1 a Scenario 2 b
Percentage crop by
Page 412
Yield Crop Value Crop Value
size grades
per value as value as
Thinning < 2 in. 2–2 in. 2–2 in. 2–2 in. > 2 in. tree per tree % of per tree % of
time (4.4 ¢ kg1) (4.4 ¢ kg1) (4.4 ¢ kg1) (13.2 ¢ kg1) (22 ¢ kg1) (kg) (US$) control (US$) control
R.E. Byers
Pink bud 0.0 0.4 9.3 38.2 52.1 9.32 1.58 100 1.54 100
FB + 1 0.0 0.5 7.3 35.3 56.9 10.95 1.92 122 1.88 119
FB + 2 0.0 1.4 18.3 50.8 29.5 12.89 1.81 115 1.70 108
FB + 3 0.1 3.0 35.5 48.3 13.1 13.02 1.43 91 1.21 76
Unthinned 3.0 19.4 46.9 27.1 3.6 14.95 1.11 70 0.65 41
Crop values per tree for scenarios 1 and 2 were added by Byers (2001, unpublished results).
a Scenario 1. Crop values: < 2 in. = 4.4 ¢ kg1; 2–2 in. = 4.4 ¢ kg1; 2–2 in.= 4.4 ¢ kg1: 2–2 in =13.2 ¢ kg1; > 2 in.= 22 ¢ kg1.
b Scenario 2. Crop values: < 2 in. = 0.0 ¢ kg1; 2–2 in. = 0.0 ¢ kg1; 2–2 in.= 0.0 ¢ kg1: 2–2 in =13.2 ¢ kg1; > 2 in.= 22 ¢ kg1.
Since cell division ceases 4–6 weeks after thinning may directly increase the fruit
bloom, fruit from trees thinned near bloom weight and elongation of ‘McIntosh’-type
are larger at harvest and have more cells cultivars (Greene et al., 1990) but not those
than fruit from trees thinned progressively of others, such as ‘Golden Delicious’, ‘Red
later after bloom (Denne, 1960, 1963; Delicious’, ‘Rome’ or ‘York Imperial’ (Greene
Sharples, 1968; Goffinet et al., 1995). The and Autio, 1994).
greater cortical cell number appears to be the
result of an increased rate of cell division
and not an extended period of cell division 16.3.3 Yields
or increased cell enlargement or intercellular
space. Even though fruit size is more corre- The economic impact of thinning on crop
lated with increased cortex cell number than value is comprised of the yield and price of
with increased cell enlargement, Sharples the most valuable size categories averaged
(1964) demonstrated that severe thinning to over 2 years or more. Even though convinc-
less than the optimum cropping 5 weeks ing data are not available, annual bearing
after bloom (after the cell-division stage) led trees within an orchard probably average
to doubling of fruit weight, due primarily to greater yields of optimum fruit sizes than
cell enlargement, cell division being only those showing a strongly biennial bearing
slightly stimulated. habit. The expectation is that a few large fruit
Seed development is essential for apple in the ‘off year’ and many small fruit in the
fruit set in most apple cultivars, but the ‘on year’ typically would result in less total
number of seeds per fruit is considered to value. Hoffman (1947) reported that when
have only a minor positive influence on fruit trees of a biennial cultivar were thinned
growth. The leaf-to-fruit ratio has by far the annually with Elgetol over a 4-year period,
greatest influence. Fletcher (1932) showed the average total yields and fruit sizes were
that hand-thinning of fruit to one fruit per 50 increased.
or 100 leaves increased fruit red colour and Parry (1974) demonstrated that removal of
size, but, when trees were fertilized, red all fruit from half of the tree was a more reli-
colour was not promoted by thinning. able method for avoiding bienniality
Preston (1954) showed substantial increases of ‘Laxton Superb’ trees than ‘whole-tree’
in yellow and red colour of ‘Duchess cluster reduction and, similarly, Preston
Favorite’ apples by thinning to 20 or 30 (1954) showed that trees that were ‘half-tree’-
leaves per fruit, but thinning to ten leaves defruited had a larger proportion of market-
per fruit only slightly affected fruit colour. able fruit over the 2-year bearing cycle.
Lawson et al. (1998) found that hand- Forshey and Elfving (1977) found that
thinning fruit to single-fruit clusters reduced ‘McIntosh’ apple yields were positively
oblique-banded leaf-roller damage to apple related to fruit numbers, but the increase in
fruit. Possible explanations were that a sin- fruit size as a result of thinning was pro-
gle larva would be more likely to damage portionally less than the decrease in fruit num-
two fruit hanging in a cluster than a single bers. Consequently, the total yield per hectare
fruit and/or that there was better pesticide of large, higher-valued fruit was either
coverage of a single fruit than of a multi- unchanged or reduced by thinning. In the year
fruit cluster. of fruit thinning, therefore, the point of dimin-
Chemical thinners or selective hand-thin- ishing returns may be quickly reached.
ning that selects for the ‘king’ fruit typically
improves the length/diameter (L/D) ratio,
colour and symmetry of the fruit and allows 16.3.4 Defruiting young trees for tree
better exposure to pest-control chemicals of growth
the remaining fruit (Westwood, 1978). Even
though fruit shape is generally not Trees grown on dwarfing rootstocks typi-
correlated with fruit density, plant growth- cally flower and fruit in the second or third
regulating chemicals (e.g. 6-BA) used for season, whereas trees of similar age grown
414 R.E. Byers
on semi-dwarf and seedling rootstocks sel- In addition, tree growth was stimulated
dom flower adequately at 4–6 years of age. more when trees were deblossomed than
Early flowering and fruiting of dwarf trees when they were defruited 36 days after
may seriously inhibit tree growth and cause bloom. Barlow (1966) demonstrated that
long-term stunting of the tree, which may thinning promoted an increase in the total
contribute to reduced tree size, poor tree number of shoots that grew longer.
structure and reduced yields for several By thinning 18-year-old ‘Sunset’/M.9
years thereafter. For this reason, defruiting of trees at progressively later times – at pink
young dwarf trees with chemical thinners or bud, FB + 7 days, FB + 14 days and FB + 21
by hand has become a routine commercial days – Quinlan and Preston (1968) recorded
practice. a 4-year incremental increase in trunk cross-
Bedford and Pickering (1916) showed that sectional area of 198%, 198%, 193% and
deblossoming young apple trees for the 155%, respectively. In addition, total bourse-
first two crop years caused trees to bear shoot length per 100 flowering clusters was
more heavily for up to 14 years after treat- greatest for the earliest thinning time and,
ment. The trees were larger and stronger over the 4-year period, was 324%, 221%,
than trees that were allowed to bear in the 151% and 149% of the control, respectively.
first two seasons. Maggs (1963) showed that Thus the increase in shoot growth per tree
deblossoming caused a threefold increase in was due to the increase in the number of
new root growth and a twofold increase in bourse shoots produced and not to their
trunk cross-sectional area when compared length. These data suggest that early thin-
with unthinned trees. Deflowering spur ning results in a greater number of fruiting
‘Rome’ trees increased terminal shoot spurs and leaves within the whole tree
growth by 52% and trunk circumference by canopy. When thinning was delayed until
47% (R.E. Byers, 1992, unpublished results). after bloom, both spur-leaf size and number
The application of gibberellins soon after were reduced (McArtney et al., 1996) and
bloom will partially, but not consistently, flower-bud size and subsequent fruit size
inhibit flower-bud formation for the subse- were decreased in the following season com-
quent season and will stimulate tree growth
pared with thinning at bloom.
in the current season independently of crop-
ping (Unrath and Whitworth, 1991).
16.4 Thinning Chemicals and Their
16.3.5 Effect of crop reduction on growth Physiological Effects
and fruiting of bearing trees
Flower thinning or flower-bud inhibition has
Thinning of bearing trees soon after bloom the advantage of maximizing fruit size,
results in greater tree growth and a greater return bloom and yield, and allows addi-
number of spurs with growing bourse tional time to reduce further as necessary the
shoots when compared with progressively crop by post-bloom chemical- or hand-
later thinning times (Preston, 1954). thinning practices. The primary disadvan-
Thinning shortly after bloom has been corre- tage of flower thinning is the potential for
lated with larger dormant buds, longer flow- subsequent damage to flowers or fruit by
ers, greater spur-leaf area, larger bourse spring frosts. Secondarily, the environmental
shoots and greater fruit size in the following conditions that have an impact on pollina-
thinning (Denne, 1963). Maggs (1963) tion, fertilization and fruit set subsequent to
demonstrated that deblossoming trees thinning are unpredictable (Batjer, 1965).
resulted in more and larger leaves, longer Flower-bud inhibition in the previous season
shoot growth, increased trunk thickening or thinning at tight cluster or pink may have
and greater root growth; however, cropping some utility for small-fruited cultivars, those
trees produced more total dry matter (vege- that flower heavily and those that are
tative growth plus crop) per unit leaf area. typically biennial bearers.
16.4.1 Pollinicides scarring and leaf injury. Pelargonic acid (57%

plus 3% related fatty acids with a total 60%
Pollinicides provide an additional physiolog- a.i.; ‘Thinex’), YI-1066 and sulphcarbamide-
ical mechanism for thinning apples that is 1-aminomethanamide dehydrogen tetraoxo-
basically different from that of hormone-type sulphate (79% a.i.; ‘Wilthin’) caused more
thinners and which could be used under cool fruit injury than mono-N,N-dimethylalkyl-
conditions where the effectiveness of hor- amine salt (Endothall 4.793% a.i., 5.5% acid
mone-type chemicals is greatly reduced. equivalent, TD 0.4EC 2337-03) or ATS (60%
Elgetol (sodium-4,6-dinitro-ortho-cresyl- ATS or 55% ATS; ‘Thinset’) in several experi-
ate) (DNOC) became the first commercially ments (Williams, 1993a; Byers, 1997; Fallahi,
important apple flower-thinning agent (Batjer 1997). Although the amount of fruit injury is
and Thompson, 1948; Williams, 1979) and usually limited, the potential for fruit scar-
remained so for over 40 years until its ring may limit the use of certain chemicals
Environmental Protection Agency (EPA) reg- for fresh-market apples. The more caustic
istration was cancelled in 1990. Hildebrand chemicals could be used, for example, where
(1944) showed that elgetol inhibited fruit set fruit are grown for processing, where surface
when applied as much as 32 h after pollina- injury would not be of economic conse-
tion, when pollen-tube growth was approxi- quence. Even though a small percentage of
mately midway down the style. fruit were marred by some of the chemicals,
Elgetol thinning at bloom was considered the resulting benefits of increased fruit size
to be a part of a total-thinning programme and return bloom may offset the negative
in the western USA that was essential for effects. Because the king flower produces the
maximizing fruit size and return bloom largest and most typically shaped fruits (for
(Williams, 1993a,b, 1995). The cancellation most cultivars), it would seem most desir-
of elgetol registration prompted the search able to apply bloom thinners in the late
for a replacement chemical (Williams, 1994), bloom to inhibit fruit set of the lateral flow-
which, based on the western US elgetol use ers and especially flowers on annual growth
pattern, could only be marginally justified in since these produce small fruit. However,
regard to new chemical registration costs. tests indicate that injury increases with later
Research has shown that elgetol, when applications and higher chemical rates
applied just before wet and humid periods, (Byers, 1997). Another disadvantage of
caused fruit and foliage injury, with erratic pollinicides is that the application window
effectiveness as a thinner. In addition, requires a short span of a few hours to make
unpredictable weather during pollination single or multiple applications. Future
and fertilization further contributed to research should focus on improved spray
unpredictable thinning related to a variable thinning chemicals that reduce or eliminate
number of flowers setting fruit. Several fruit russet and leaf injury in single or
chemicals that were effective as peach thin- multiple applications, since each chemical
ners were found to be effective apple-bloom may have differences in residual half-life and
thinners (Endothall, ammonium thiosul- cumulative phytotoxicity when used in
phate (ATS) and long-chained fatty acids) multiple applications.
(Byers and Lyons, 1985; Byers et al., 1985a,b; The search for pollination/fertilization
Williams, 1993a, 1995; Southwick et al., 1996; inhibitors that do not affect fruit finish
Bound and Jones, 1997; Byers, 1997; Fallahi, deserves considerable attention for fruit des-
1997). The use of ATS for flower thinning has tined for the fresh market. Certain fungicides
been commercially practised in the USA for such as Captan and Triforine (Funginex) are
over 10 years and is the standard material strong inhibitors of pollen-tube growth in
used in much of Europe. vitro (Fell et al., 1983). These chemicals are
Because the apple receptacle and spur also known for their non-phytotoxic effects
leaves are exposed to spray chemicals before, to fruit and foliage. There has been specula-
during and after flower opening, caustic tion that fungicides that inhibit pollen-tube
bloom-thinning sprays may cause some fruit growth in vitro might reduce fruit set in the
416 R.E. Byers
field if applied at bloom (Fell et al., 1983; for inhibition of flower-bud formation
Gomma, 1989). Preliminary data indicate (Dennis and Edgerton, 1966; Tromp, 1982;
that Captan reduced fruit set when applied McArtney and Li, 1998). Tromp (1982) found
with a handgun during bloom but not when that GA4+7 more effectively reduced flower-
applied with an air-blast sprayer (R.E. Byers, ing than did GA3 on both spurs and 1-year-
1993, unpublished results). Since fruit pro- old shoots. However, Marino and Greene
duced by late bloom (flowers borne laterally (1981) found that GA3 was more effective on
on the previous season’s terminal growth) 1-year-old shoots and GA4+7 was more effec-
are frequently smaller than those produced tive on spurs at decreasing flower-bud for-
by flowers borne on spurs, fungicides mation. Applications of gibberellins must be
applied at early petal fall could be beneficial made at bloom or shortly thereafter to be
for eliminating fruit set of late flowers, effective on spurs, but applications up to 60
thereby improving fruit size. Screening polli- days AFB are effective on 1-year-old shoots
nation inhibitors for use on late-opening (Tromp, 1982). Gibberellins used to reduce
flowers and that do not cause russeting of russeting of ‘Golden Delicious’ fruit have
earlier-formed fruit is important. also shown some inhibition of flower-bud
At bloom, the percentage of flowers open formation (Meador and Taylor, 1987; Greene,
between adjacent trees in the same row may 1993). It is believed that gibberellins interfere
range from 40 to 95%. In addition within the with the early phases of bud primordia
same block, terrain, microclimates, cultivar, development long before flower buds are
strain, and soils can also affect flower open- microscopically visible. Although less effec-
ing, pollen tube growth rate, natural fruit set; tive for flower-bud inhibition, GA3 may be a
and thus, if a pollination inhibitor is applied better choice economically, since the price of
to all the trees in the block at the same time, GA4+7 may be five times that of GA3.
the potential exists for creating wider differ- To maximize tree growth, Unrath and
ences in fruit set than existed naturally. Whitworth (1991) attempted to completely
Multiple applications applied at intervals of inhibit flowering of young non-bearing ‘Red
2–3 days may reduce thinning variation Chief Delicious’ trees. In one experiment,
caused by differences in flower opening, but multiple applications GA4+7 at rates of 250
test results have not been consistent (Byers, mg l1 or 500 mg l1 reduced flowering by
1997). 95% and 99%, respectively. In two other
experiments, similar treatments gave very
little or no suppression of return bloom;
16.4.2 Gibberellin for inhibition of flower- however, the timing of GA4+7 applications
bud formation may have been too late.
In bearing ‘York Imperial’/MM.111 trees,
Since heavy flowering in one year inhibits a single spray of GA3 in the ‘off year’ pro-
the growth and development of the bourse vided more regular cropping for 4 subse-
flowers in the subsequent season, partial quent years (Byers et al., 2000b).
inhibition of flowering with gibberellin
sprays may promote fewer and larger flow-
ers. McArtney (1994) demonstrated that a 16.4.3 Hormone-type plant-growth
single spray of gibberellin A3 (GA3) or GA4+7 regulators (PGR) for bloom and
at full bloom in the ‘off year’ reduced the fruit thinning
subsequent season’s flowering in the ‘on
year’ and the severity of biennial bearing of Because hormone-type thinners are less
‘Braeburn’ trees. Increasing concentrations effective during bloom, they have been
of a gibberellin spray linearly decreased primarily used for fruit thinning, in contrast
flowering the following year and promoted to pollination inhibitors, which have been
flowering 2 years after application. Several used for flower thinning (Williams, 1979;
experiments have shown GA7 to be more Wertheim, 1997; Dennis, 1986; Dennis,
effective than GA3 and that GA4 is ineffective 2000). Hormone thinners are not dependent
on flower opening and are less injurious to uptake under increasing temperature and
fruit or foliage than are many caustic decreasing humidity conditions throughout
pollinicides. Since insecticides such as car- the day has opposing influences, which
baryl, dylox, morestan and oxamyl kill wild appear to compensate for one another.
or domesticated pollinating insects, certain The fundamental mechanism(s) by which
hormone-type thinner combinations should a hormone thinner causes fruit abscission
not be applied during bloom. The value of is not known, but Ebert and Bangerth
very early thinning with chemicals such as (1982) suggested that a major factor in the
NAA, 2-chloroethylphosphonic acid (ethep- mode of action of carbaryl, ethephon and
hon), carbaryl (XLR formulation), 6-BA or naphthaleneacetamide (NAAm) is a reduc-
Accel at petal fall is substantial (Jones et al., tion in indole-3-acetic acid (IAA) transport.
1989, 1990, 1991; Williams, 1993b; R.E. Williams (1981) found that NAA and car-
Byers, 1994–1999, unpublished results). baryl caused elevated ethylene levels in
Pollinicides, however, provide an alterna- apple fruits similarly, but NAA caused a
tive to hormone thinners when the effec- greater stimulation of ethylene in leaves.
tiveness of hormone thinners is limited due Aminoethoxyvinylglycine hydrochloride (N-
to low-temperature conditions. (phenylmethyl)-1H-purin-6-amine, or AVG),
Chemical thinning has typically been an inhibitor of ethylene biosynthesis, applied
quite variable, depending on factors such as before or after either NAA or carbaryl sup-
the orchard, cultivar, location and year. pressed ethylene and abscission of fruit, but,
Although 50 possible factors might influ- when either ethephon or ethylene gas was
ence the effectiveness of NAA thinning, applied to the trees, fruit abscission was not
most are non-significant (Hennerty and inhibited. Ward et al. (1999) recently found
Forshey, 1972). Of all these factors, the most that cellulase activity did not appear until
significant was a positive relationship approximately 4 days after the application of
between the number of fruits removed and carbaryl + ethephon, but fruit diameter,
the number of fruits naturally set on the water potential and starch were reduced
tree. At the spray date, fruit set was posi- within 2 days after treatment. Since low light
tively related to the nitrogen content of the further stimulates fruit abscission, the role of
spur at full bloom and to shoot vigour at photosynthates for fruit retention may be
green tip, but negatively related to spur fundamental hormone to the action of chem-
vigour. Fruit thinning was not related ical thinners (Byers et al., 1991).
directly to tree vigour, soluble or total nitro-
gen or soluble, reserve or total carbohy-
16.4.3.1 Naphthaleneacetic acid
drates. The physiological tree ‘condition’
was thought to have a greater effect on flow- In 1941, auxin-type sprays (NAA and NAD)
ering and fruit set than the overall response were found to cause fruit abscission
of the tree to thinning chemicals. (Burkholder and McCown, 1941), but their
Foliar penetration and movement of hor- commercial value was not apparent until the
mone-type thinners to their site(s) of action early 1950s. Cultivars that reached maturity
are required for fruit thinning to be achieved. during early, mid- and late summer were
The cuticle on the under leaf surface permits typically thinned by the more mild NAD,
approximately tenfold greater NAA pene- because NAA caused foliage injury, inhibi-
tration than the upper cuticle (Norris and tion of fruit growth, premature ripening and
Bukovac, 1969). Penetration of NAA or fruit splitting. In contrast, cultivars that
naphthalene acidimide (NAD) is highly mature in the autumn were typically thinned
dependent on temperature, especially with NAA (Batjer, 1965), since many of these
between 15 and 25ºC, and is also positively detrimental effects were not observed.
correlated with increasing humidity. D.W. NAA alone has not caused specific fruit-
Greene (1998, personal communication), finish or shape problems. To reduce the neg-
however, has pointed out that 14C-NAA ative effects of NAA on fruit growth, lower
418 R.E. Byers
rates of NAA (2–5 mg l1) in combinations followed the negative effect of NAA on
with carbaryl, oxymal and/or adjuvants fruit reducing sugars; thus, ethylene was
have given excellent thinning in most years. not thought to be the cause of NAA-induced
Combinations of NAA and carbaryl are abscission. However, Curry (1991) demon-
widely used for chemical thinning of many strated that NAA promoted ethylene
apple cultivars, but NAA may cause serious evolution very soon after application.
overthinning and/or dwarf fruit in certain He proposed that 1-aminocyclopropane-1-
cultivars (Rogers and Williams, 1977). carbonate (ACC) produced in the leaves may
Several papers and reviews suggest that be transported to the abscission zone, where
many cultural and environmental factors it is converted to ethylene and causes fruit
influence NAA thinning, and local and abscission.
regional conditions are important to deter- Stopar et al. (1997) showed that NAA
mining recommendations (Westwood, 1978; suppressed photosynthesis by 10–24% over
Dennis, 1979; Lehman et al., 1987; Byers et al., a 15-day period and was concentration-
dependent. NAA sprays have been shown
1990b). NAA and NAD alone or in various
to reduce stomatal opening (Snaith and
combinations with gibberellins were found
Mansfield, 1984) and result in less reducing
to increase the incidence of dwarfed fruit
sugars and sorbitol in fruit (Schneider and
(pygmy fruit) and retard fruit growth in
Lasheen, 1973; Schneider 1975, 1977). Since
most spur ‘Delicious’ and ‘Fuji’ strains
shading trees for a 2–3 day period caused
(Henderson and McBurnie, 1943; Rogers and substantial fruit drop (Schneider, 1978; Byers
Williams, 1977; Byers, 1978; Westwood, 1978; et al., 1990a,b), reduced photosynthesis is
Byers et al., 1982). known to cause fruit thinning. A long period
Soon after application to leaves, 14C-NAA of reduced photosynthesis caused by NAA
was found in apple fruits and seeds (Stopar et al., 1997), along with low environ-
(Williams and Batjer, 1964). Luckwill (1953) mental light conditions, strongly supports
suggested that seed abortion may be respon- the view that fruit abscission involves
sible for fruit abscission. However, Luckwill energy-driven processes that operate over a
and Lloyd-Jones (1962) recovered only 0.2% 2- or 3-day period.
of 14C-NAA applied to apple leaves from
seeds after an interval of 5 days, and all of
the applied NAA was metabolized. This 16.4.3.2 Carbaryl
suggests that fruit abscission is not due to In 1958, carbaryl was found to be a mild
the direct toxic action of NAA to seeds. Four thinner (Batjer and Westwood, 1960) and
days after placing NAA on the upper leaf was rate-insensitive above 750 mg l1
surface, 80% of the NAA was lost (presum- (Southwick et al., 1964; Way, 1967). In addi-
ably to ultraviolet (UV) light destruction), tion, multiple applications typically did not
10% was found within the leaves and 10% increase thinning (Byers and Carbaugh,
was still on the leaf surface. This neutral 1991). Most spur ‘Delicious’ strains normally
metabolite in the leaf was devoid of auxin- set heavy crops, and carbaryl alone was
like activity, but, since non-auxin compounds found to be inadequate for thinning
also cause fruit thinning, it is possible that it (Henderson and McBurnie, 1943; Rogers and
had thinning activity. In addition, since NAA Williams, 1977; Byers and Carbaugh, 1991).
has been shown to thin both seeded and The effectiveness of carbaryl was increased
seedless fruit of one cultivar (Dennis, 1970), by the addition of superior spray oil (Byers,
the seed-abortion theory is not likely to be 1978; Byers et al., 1982), possibly because car-
the mode of thinning action of NAA. baryl is more soluble in oil than in water, but
NAA has been shown to cause an increase oil is also known to inhibit photosynthesis
in fruit ethylene production (Walsh et al., midly. Carbaryl is not thought to affect
1979). The leaf epinasty response caused by return bloom directly.
NAA is a typical ethylene response, but, in Under certain undetermined environmen-
Schneider’s (1975) work, ethylene evolution tal conditions, carbaryl may cause fruit
injuries, characterized by lenticle spotting. If 16.4.3.3 Oxamyl

there is substantial injury, the fruit may be
Vydate, a systemic carbamate, at its highest
scarred and misshapen on the lower side
insecticidal rate was found to be a mild thin-
and/or calyx end, particularly with spur
‘Delicious’ (R.E. Byers, 1981, unpublished ner similar to carbaryl (Byers et al., 1982).
results; Byers et al., 1982, 2000a; R.C. Unrath, Oxamyl used as a thinner did not affect mite
1998, unpublished results). High rates or populations, but did increase side-russet
multiple applications of carbaryl plus adju- development in ‘Golden Delicious’ fruit
vants (oil or surfactants) or combinations and in other cultivars prone to this problem;
with other fruit thinners, such as carbaryl furthermore, lenticle enlargement may also
plus Accel (6-BA + GA4+7) plus oil, may occur (Byers et al., 1982). Certain combina-
cause additional injury. Artificially shading tions, such as Oxamyl + Accel + oil, may
trees on the day of the carbaryl spray or cause substantial red-colour inhibition of
every third day substantially increased injury. spur ‘Delicious’ fruit. Oxamyl alone under
At application time, small fruit (10 mm) seem certain environmental conditions (not yet
to be more susceptible to injury than larger defined) may inhibit red colour between
fruit (17 mm) sprayed later in the spring. The lenticles and cause a red-spot appearance
injury may be severe in some years, but not around the lenticles on the fruit surface (R.E.
others (Byers et al., 2000a). The cause for year- Byers, 1995, unpublished results).
to-year injury variation has not yet been
determined (Plate 16.1).
16.4.3.4 Ethephon
Williams and Batjer (1964) found that fruit
were more susceptible to abscission if only Ethephon is considered a very dose- and
fruit were treated than if only leaves were temperature-dependent chemical thinner
treated. 14C-carbaryl applied to fruit moved (Koen and Jones, 1985) and is the only mater-
into the vascular tissues of the fruit (the pro- ial that can effectively thin from bloom
posed site of action). Knight (1983) believed through to a stage when fruit are larger than
that the bourse-shoot leaves could absorb about 25–30 mm in diameter (Veinbrants and
and translocate sufficient carbaryl to cause Hutchinson, 1976). It is typically used when
fruit thinning and this proposition was fur- earlier chemical thinning sprays have failed
ther supported by Byers et al. (1991), who to thin adequately. Considerable experience
found that treating either leaves or fruit is needed to use this chemical most effec-
caused fruit thinning. tively, since ethephon can be erratic and can
Carbaryl is widely known to kill preda- trigger total fruit drop. Information indicates
tors of phytophagous mites and the use of that the volume of water and method of
carbaryl frequently causes mite populations application may have more influence on
to fluctuate out of control (Byers et al., 1982). thinning than high temperature or chemical
Its use, therefore, presents difficulty in main- concentration (Unrath, 1978; Jones et al.,
taining integrated pest management (IPM) 1990, 1991). In Virginia, spur ‘Delicious’,
programmes (Hislop and Prokopy, 1981; ‘York Imperial’, ‘Gala’ and ‘Winesap’ are less
Elfving and Cline, 1993a), particularly if two responsive to ethephon, while ‘Golden
applications of carbaryl are made. The Delicious’ and ‘Rome’ are more easily over-
carbaryl formulation Sevin XLR has been thinned than most other cultivars (R.E.
found to be less injurious to mite predators Byers, personal observations). Lower rates of
and bees and thus has been successfully ethephon and/or lower volumes of water
used at petal fall for thinning. The addition may be required for these cultivars.
of low rates of a 70 s, delayed dormant, Combinations of ethephon and carbaryl have
superior oil (0.25–0.5%) to carbaryl thinning given good thinning of spur ‘Delicious’ in
sprays has provided some initial suppres- most Virginia and North Carolina tests
sion of mite populations (R.L. Horsburgh (Unrath, 1978; Byers et al., 1990a; R.P. Marini,
and L. Cobb, 1989, unpublished results; 1996, unpublished results). Since thinning
Biggs et al., 1999). apples with ethephon can be erratic, registra-
420 R.E. Byers
tion and recommendations for using ethep- Carbaugh, 1991; Greene and Autio, 1994).
hon for thinning need careful consideration The increased fruit size caused by 6-BA has
and coordination between the chemical com- been attributed primarily to a direct stimula-
pany, state and federal registration agencies. tion of cell number in the fruit cortex during
Inhibition of fruit growth has been of the cell-division stage and not to cell enlarge-
concern in several ethephon experiments ment (Wismer et al., 1995). 6-BA also has
(Edgerton and Greenhalgh, 1969; Way, 1971; been shown to increase L/D ratio in some
Chiba et al., 1980; Knight, 1980; Ebert and cultivars, but may have no effect in others
Bender, 1986), but not in others (Veinbrants (Jones et al., 1997).
and Hutchnson, 1976; R.E. Byers, Stopar et al. (1997) found that 6-BA had no
1980–2000, unpublished results). In effect on photosynthesis but Yvan and
Tasmania, ethephon thinning sprays at Greene (2000) found that 6-BA-treated leaves
bloom were found to reduce the L/D ratio, and fruit had higher night respiration rates,
but addition of 6-BA (Cytolin) restored fruit thus reducing the total photosynthates avail-
shape (Bound et al., 1993). able. Although 6-BA sprays have been
Ketchie and Williams (1970) found that shown to increase ethylene evolution from
ethephon (250–1000 mg l1) applied in the fruit before abscission (Kondo and Mizuno,
1–2-month period prior to autumn leaf drop 1989), the magnitude of the ethylene increase
dramatically reduced fruit set and vegetative- was not considered large enough to be the
shoot length the next spring. This property primary cause for thinning (Greene et al.,
might be used to inhibit flowering if applied 1992). Unlike NAA, application of 6-BA to
after harvest in the ‘off year’ of the biennial the fruit alone caused modest fruit thinning,
bearing cycle. The potential for producing but the maximum effectiveness was obtained
smaller fruit exists but the early flower-bud by application to both leaves and fruit
inhibition may compensate for the direct (Greene et al., 1992).
effect of ethephon on fruit size. Accel, a commercial formulation contain-
ing a combination of 6-BA and a low
concentration of GA4+7, was registered for
16.4.3.5 Dylox
thinning apples for the first time in 1995 in the
The Chinese reported that the insecticide USA. Since gibberellins are known to promote
Dylox thinned apple fruit and increased fruit retention, the low gibberellin levels in the
ethylene evolution of the fruit (Shen and Accel formulation may reduce the thinning
Sen, 1985). In Virginia, Dylox was found to response (T.L. Robinson, 1999, unpublished
thin spur ‘Delicious’ with no detrimental results). Cultivars naturally prone to pygmy
side-effects (R.E. Byers, 1991, unpublished fruit development should not be sprayed with
results). This insecticide could be very useful gibberellins, since a greater number are set
to the apple industry since it has only a (Byers and Carbaugh, 1991; Greene and Autio,
minor impact on mite predators and yet it is 1994). In certain cultivars (‘York Imperial’),
toxic to leafhoppers, leaf-miners and several Accel may increase the number of seedless
other apple pests. fruit, even though they may achieve full size
(R.E. Byers, 1995, unpublished results).
6-BA combined with carbaryl was found
16.4.3.6 6-Benzyladenine
to be more effective than either compound
The cytokinin 6-BA has been shown to be an alone and is considered to be one of the most
effective fruit thinner and has increased fruit effective chemical thinning combinations
size beyond the thinning effect for (Byers and Carbaugh, 1991). The addition of
‘McIntosh’ types (McLaughlin and Greene, superior oil to 6-BA or its combination with
1984; Greene et al., 1990, 1992; Byers and carbaryl substantially increases effectiveness
Carbaugh, 1991; Elfving and Cline, 1993a,b; (Byers and Carbaugh, 1991; Greene et al.,
Greene and Autio, 1994; Wismer et al., 1995), 1992). Combinations of 6-BA and carbaryl
but not for other cultivars, such as ‘Golden may cause additional fruit russet of ‘Fuji’
Delicious’ and ‘Red Delicious’ (Byers and and ‘Golden Delicious’ (Bound et al., 1991a,
1993) and, when 6-BA or Accel is combined 16.3.3.8 Other chemicals that may
with Oxamyl plus oil or surfactants, fruit influence fruit set
russet in the lenticels and fruit colour may
Season-long applications of sterol-inhibiting
be affected (R.E. Byers, 1995, unpublished
fungicides (bitertanol, etaconazol or fenari-
results). Trees should not be sprayed with
mol) have caused significantly less return
combinations of 6-BA and NAA, since a sig-
bloom when compared with a standard fungi-
nificant number of seedless pygmy fruit may
cide programme (Latham et al., 1985).
occur (Bound et al., 1991b).
Triadinefon was found to affect fruit size, fruit
shape and fruit set (Strydom and Honeyborne,
16.4.3.7 Photosynthetic (PN) inhibitors 1981). Since sterol inhibitors and plant-growth
Since short periods of artificial shade may retardants (Miller, 1988) have been shown to
cause fruit abscission, photosynthetic inhibi- increase fruit set and affect fruit size and
tors were screened for their effectiveness shape, they should be evaluated for interfer-
as thinning chemicals and for injury to ence with chemical-thinner efficacy.
fruit and leaves (Byers et al., 1984, 1985b, Prohexadione-Ca (Apogee®), a new plant-
1990a,b). Extremely low rates of herbicides growth retardant registered for apples, has
have been found to cause fruit abscission been shown to promote additional fruit set
without effects on the remaining fruit or alone (D.W. Greene, 1997, unpublished
leaves, but others may cause unacceptable results), but the fruit appear to respond to
leaf injury (Byers et al., 1990a,b). Terbicil was various chemical-thinner combinations in a
one of the more promising chemicals, but its similar way to non-treated fruit (Byers et al.,
persistence and leaf injury in some cultivars 2000a).
(e.g. ‘Golden Delicious’) and the lack of The potential exists for the use of specific
interest from the chemical company for reg- pesticides (superior spray oil, liquid lime sul-
istration have stopped its development as a phur, organic phosphates, sterol inhibiting
thinning compound. Most photosynthetic fungicides) or plant growth regulators (ethep-
inhibitors available are selected for their hon) prior to, or in conjunction with, thinning
persistence for use as herbicides for killing chemicals to pre-condition trees to make them
weeds, but short-lived inhibitors, either more responsive to thinning when hormone-
alone or as an additive to other registered type chemicals are applied. This strategy
chemical thinners, could be a more useful could be very useful for difficult-to-thin culti-
tool, with less injury to leaves. vars (such as ‘Fuji’, ‘Gala’, ‘Golden Delicious’,
Certain pesticides registered for use on ‘Pink Lady®’) and/or regions where high light
apples reduce photosynthesis of apple leaves or cool conditions make for difficult thinning.
by 20–30% following multiple applications in
the laboratory, such as superior oil (Ayers and
Barden, 1975; Ferree and Hall, 1975; Ferree et 16.5 Adjuvants
al., 1976; Wood and Payne, 1984), ethion
(Heinicke and Foott, 1966), dicofol (Sharma et The use of a suitable surfactant has been
al., 1977), fenvalerate and diazinon (Heinicke thought to reduce thinning variability
and Foott, 1966) and other organic phos- caused by widely different environmental
phates (Pickett et al., 1952; Ayers and Barden, conditions (Westwood and Batjer, 1960). In
1975; Ferree and Hall, 1978; Anderson, et al., several regions where adjuvants are com-
1986). Several of these materials in combina- mercially used, NAA is typically reduced to
tion with carbaryl potentiate thinning (Byers one-half the recommended rate, resulting in
and Carbaugh, 1991). Since trees sprayed with a substantial cost savings. The use of a
carbaryl in field experiments have not shown surfactant with NAA is general practice in
reductions in photosynthesis, inhibition of some areas (Williams, 1979), but not in others
photosynthesis was not considered the pri- (Robinson et al., 1998). There is evidence to
mary cause of fruit abscission (R.E. Byers, indicate that the chemistry of each thinning
1988, unpublished results). compound interacts specifically with each
422 R.E. Byers
adjuvant, such that each combination more uniform thinning results by TRV cali-
requires testing for its effectiveness. bration with ethephon, NAA plus carbaryl
Several investigators have found that and NAA plus ethephon than by using a
surfactants, superior oil (Byers, 1978; Ebert specific water and chemical rate per hectare,
and Bender, 1986), penetrants, fertilizers regardless of tree size. TRV calibration of
(Horsfall and Moore, 1961) and pesticides may air-blast sprayers was an attempt to maintain
increase thinning to varying degrees. Greene an equivalent deposit of chemical and/or
and Bukovac (1971) have shown that surfac- water volume per unit leaf area when tree
tants may significantly decrease surface ten- sizes changed (Byers et al., 1971, 1984, 1989;
sion and increase absorption, but that Herrera-Aguirre and Unrath 1980; Sutton
absorption was not directly related to surface and Unrath, 1988 a,b; Byers, 1989).
tension. Certain adjuvants alone or in combi- Several studies have shown that low
nation with other chemicals may increase thin- water-volume sprays were less effective for
ning by stimulating ethylene production. Oils, thinning than were high water volumes
liquid lime sulphur and/or organic phos- (Rogers and Thompson, 1969; Rogers and
phates may reduce photosynthesis, respiration Williams, 1977; Jones et al., 1988, 1991; R.P.
or some other metabolic mechanism(s). Marini, Virginia, 1995, personal communica-
Greene and Bukovac (1971), using pear- tion). In one experiment, Jones et al. (1991)
leaf discs in the laboratory, established that demonstrated that ethephon effectiveness
surfactants enhanced penetration of NAD applied with an air-blast sprayer was posi-
through the lower leaf surface, but not tively related (linearly) to the water volume;
the upper. The lower leaf surface absorbed however, increased chemical rates per hectare
15-fold more NAD than the upper surface did not increase thinning. Even though
after 48 h. Penetration was not affected by ethephon thinning tests in Virginia have also
pH, but the pKa for NAD was 13; thus the been inconsistent, the chemical rate appeared
NAA molecule was non-dissociated over the to be more important than the water rate per
pH range studied (pH 3–7 range). ha in the range of 935 l ha1 to 3740 l ha1.
Superior oil and silicone surfactants Looney and McKellar (1984) found that
appear to be among the best of the penetrants spray volumes from 560 to 4400 l ha1 did
used for chemical thinners, but in some tests not have a major influence on thinning, but
these compounds increased side and/or better thinning was achieved with reduced
lenticel russet or fruit colour if the chemical spray volumes in two of three experiments.
thinner was prone to producing injury (R.E. Additional experiments have shown that
Byers, 1995, unpublished results). low-volume (LV) or ultra-low-volume (ULV)
sprays for apple thinning can be as effective
as high-volume (HV) sprays (Oakford et al.,
16.6 Application Considerations 1991, 1994, 1995). However, the success of
LV or ULV sprays is dependent on control-
When chemical thinners were first tested, ling droplet size by using air-shear machines
most were applied with a handgun sprayer that produce droplets in the 60–120 µm size
at high water volumes to the point of drip range. Most HV sprayers produce droplet
from foliage, and the amounts of chemical sizes greater than 150 µm, which more easily
and water were, therefore, frequently not run off the leaf surface. In addition, the high
known. Air-blast spray applications were pressure used in many HV sprayers pro-
frequently not as effective as handgun appli- duces a considerable number of droplets
cations. Sutton and Unrath (1988a) demon- that are less than 50 µm and this size does
strated that handgun applications increased not impinge on the target and the droplets
chemical deposits by 60% or more over air- drift significantly. Spraying time where LV
blast tree-row volume (TRV) spray rates and sprayers are used can be reduced by as
that fruit thinning and disease control were much as 60% over that of HV spraying, and,
more effective using handgun applications. according to Oakford et al. (1995), chemical
Herrera-Aguirre and Unrath (1980) found rates may be reduced to some extent. Bound
et al. (1997) increased the chemical-thinning 16.6.2 Effect of application temperature on

effectiveness of conventional air-blast chemical fruit thinning
sprayers by using a redesigned nozzle that
produced finer droplets at low volumes. Previous reviews suggest that chemical
These nozzles (developed by Delavan-Delta thinning increases with higher tempera-
Inc.) produced a greater number of optimum- tures (Westwood, 1978; Williams, 1979).
sized droplets than the traditional HV Jones and Koen (1985) showed a marked
hydraulic-sprayer nozzles. increase in fruit thinning by ethephon held
In controlled laboratory spray studies, in controlled-environment rooms as tem-
Knoche et al. (1998) showed that either peratures increased from 4 to 24°C. These
decreasing droplet size and/or increasing trees were exposed to the specified temper-
carrier volume (at lower volumes) increased atures 4 h prior to, during and after and
biological performance of 2,4-dichloro- held for 24 h.
phenoxyacetic acid (2,4-D) and daminozide In a laboratory experiment using pear
at a constant dose, but had little effect on the cuticles, Greene and Bukovac (1971) showed
performance of GA3. These data demonstrate substantial increases in penetration of NAD
that there is an optimum spray volume and at increased temperatures. Edgerton and
Haeseler (1959) reported similar results from
droplet size that can maximize the biological
NAA in a greenhouse study. Observations
activity of some growth regulators, but not
that thinning increased with higher field
others.
temperatures have been explained by greater
chemical absorption at higher temperatures.
16.6.1 Drying rate D.W. Greene (Massachusetts, 1998, personal
(dew/rain/humidity/rewetting) communication) has found little evidence for
increased absorption of NAA with increased
temperatures in the field, presumably
While a droplet is drying, there is an increas-
because drying time decreased with lower
ing rate of chemical concentration and an
humidity as temperatures increased. Jones et
immediate uptake of the chemical, at least
al. (1988) found no relationship between tem-
under laboratory conditions (Greene and
perature and humidity with fruit thinning,
Bukovac, 1971). When chemical solutions are but they found a linear decrease in thinning
almost dry, uptake of the remaining chemi- by NAA from 0800 to 2000 h. It is possible
cals slows dramatically. If a surfactant is that the combined effect of a dark (night)
used, over 50% of NAA may be absorbed period followed by a chemical thinning
during the drying process, but the remaining spray may explain the greater effect of the
chemical may be destroyed by light in 1–2 early-morning spray timing.
days. The hydration properties and/or When NAA is used on most cultivars at
decrease in pH of ammonium nitrate solu- cool temperatures, pygmy fruit that are in the
tions with NAA may increase diffusion by 10–20 mm fruit size often remain on the tree,
increasing the number and mobility of the shrivel and die in the dormant season. These
non-dissociated NAA species (Fader and dead fruit frequently harbour rot organisms
Bukovac, 2001). Since carbaryl is not and increase innoculum the following season
destroyed by light, it may remain unchanged (K.S. Yoder and R.E. Byers, 1996, unpublished
on the leaf surface for a week or more, and results). Though very controversial, early
rewetting by dew or a light rain may facili- work indicated that temperatures of 13–29°C
tate additional absorption. at the time of application did not affect thin-
Ethephon is pH-dependent and, as the ning by NAA or NAD. In addition, when
droplet dries, the pH may increase or temperatures were cool, differences in chemi-
decrease, causing differences in evolution and cal rate of 50% frequently gave no difference
absorption. After ethephon application, leaf in thinning response (Thompson, 1957).
ethylene evolution is very temperature-depen- Little information exists on application
dent and may be continuously evolved for temperature and its influence on thinning
more than a week at 21°C (Byers et al., 2000b). with hormone-type chemicals. In one field
424 R.E. Byers
experiment, applications of carbaryl at 2 h ceptible to artificial or environmental fruit

intervals from 0600 to 2000 h to ‘Empire’/ losses by shading (Byers et al., 1991).
Mark trees caused similar thinning, even
though application temperatures ranged
from 18 to 36°C (R.E. Byers, 1996, unpub- 16.7.1 Effect of light and darkness on apple
lished results). In another field experiment, fruit retention and thinning
‘Starkrimson Delicious’/Mark trees sprayed
with ethephon, Accell or carbaryl at 0600, Apple trees grown in the eastern USA are typ-
1400 or 2000 h caused similar thinning, even ically exposed to 2–3-day periods of low light
though temperatures were 14, 34 or 20°C, during cloudy/rainy periods. The low-light
respectively; thus no strong relationship of conditions can reduce photosynthetically
temperature at application time with fruit active radiation (PAR) to 10–15% of full sun.
thinning has been reported. Artificially shading trees (92% polypropylene
shade material) for 2–3 consecutive days in the
period of 14–28 days after bloom was found to
16.7 Influence of Photosynthetic Reserves reduce fruit set of spur ‘Delicious’ apple trees
on Fruit Set and Chemical Thinning of (Fig. 16.1). Under these shade conditions, the
Apples Following Ovule Fertilization smallest fruit stopped growing before the
larger fruit (Byers et al., 1991; Fig. 16.2). In one
In the spring when the temperatures have test, if 1–2 days of sunlight separated each
increased, photosynthetic reserves stored period of 2–3 days of artificial shade, much
from the previous season are used to promote less fruit abscission occurred (Table 16.2). In
the first flush of growth and to retain fruit addition, natural ‘June’ fruit drop appeared to
(Heinicke and Childers, 1937; Abbott, 1960; be related to weather events that produced 2–3
Hansen, 1971). During the period from bud days of intense cloudy weather at average
break to approximately 30 days AFB, there is temperatures above 16.7°C (Fig. 16.3; R.E. Byers
a net loss of organic carbon reserves. The et al., 1991, 1998, unpublished results).
greatest net deficit occurs about 20 days after If we speculate that 10% PAR is required to
flowering when fruit are about 12 mm in fruit maintain tree and fruit respiration during the
diameter. This photosynthetic deficit corre- daylight hours (the period from the hour after
sponds to the time when trees are most sus- sunrise to the hour before sunset), the net
30 80
% Fruit that dropped that reached
Non-shade
Shaded (24–26 May) 70
maximum size on each date
25 Sunlight (PPF)
60
PPF (mol m–2 day–1)
20
50
15 40
30
10
Shaded 20
5
10
0 0
24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47
Days after full bloom (1988)
Fig. 16.1. ‘Redchief Delicious’ fruit diameter and daily photosynthetic photon flux (PPF) during May and June
1987. Shading trees for 3 days caused a high percentage of fruit to stop their growth on the 1st to 4th day after
the period. A few larger fruit continued to grow for up to 8 days before stopping growth and later abscising.
✩Statistical differences P = 0.05, for six trees (50 tagged fruit per tree measured daily). (From Byers et al., 1991.)
18 Growth curves for

fruit stopping growth
17 each date 24% 75% ‘King’
11 May Fruit set Fruit set
16
12 May
15 13 May
14 May
14 15 May
Fruit diameter (mm)
16 May
13 17 May
19 May
12 Set fruit
Percentage Percentage
11 Shaded all fruit drop ‘King’ drop
10
13% 74%
9
8 24% 70%
7 27% 37%
6
12% 18%
5
9 10 11 12 13 14 15 16 17 18 19 20 21
Days after full bloom (May 1988)
Fig. 16.2. Shading ‘Redchief Delicious’ trees for 3 days, 9 days AFB, caused 25% fruit drop of ‘king’ fruit
and 75% of ‘side’ fruit to drop. The largest fruit destined to drop remained on the trees for more than 7 days
after the end of the 3 days of shade (drop-induction period). The smallest fruit that dropped reached their
maximum size during and soon after the shade event and remained on the tree for 7 days or more before
abscising. (From R.E. Byers, 1988, unpublished results.)
Table 16.2. Effect of shade (92%) intervals on ‘Campbell Redchief Delicious’/MM.111 apple tree fruit
abscission (from Byers et al., 1991).
Shaded days Fruit per cm2

(days AFB) cross-sectional
area of trunk
Treatment1 30 31 32 33 34 35 36 (+ 56 days AFB)
Control 6.96 a2
Shade XX XX XX 2.40 c
Shade + sun XX XX XX 4.29 b
Shade + sun X
X XX 3.00 bc
Shade + sun X
X XX 4.51 b
Shade + sun X
X XX 6.11 a
Natural light levels 55 35 46 53 66 65 62
(PPF mol m2 day1)
1 Full
bloom occurred 22 April 1988. Shade treatments were started 31 days AFB when fruit were
20.3 mm ± 0.37 mm in diameter.
2 Means of six trees. The control mean represented 857 fruit. Means separation within columns by
Duncan’s multiple range test, 5% level.

PPF, photosynthetic photon flux.
426 R.E. Byers
Fruit diameter (mm)

40 Diameter of dropping fruit 70

35
Fruit diameter (mm)
60
30 50
25
40
20
30
15
10 20
5 10
0 0
1 6 11 16 21 26 31 36 41 46 51 56
Days after full bloom (1987)
Fig. 16.3. ‘Redchief Delicious’ fruit diameter and daily photosynthetic photon flux (PPF) during May and
June 1987. Fruit drop (‘June drop’) was initiated by naturally occurring low-light period(s). Near the end of a
third major cloudy period (22–25 days AFB), average fruit diameter was 22–23 mm. Fruit dropping 38 days
AFB were 21.4 mm ± 0.3 mm (after reaching their maximum size, these fruit probably shrank by 1–2 mm).
(From Byers et al., 1991.)
photosynthetic needs of the tree may not be mately 10–15 mm diameter) were placed in
adequately met for the 24 h period. At night, controlled-environment rooms for 2–6 days
no light-energy input occurs and all respiration in darkness. In one experiment, trees held at
is dependent on photosynthetic reserves accu- 4.4°C in darkness for 3 days did not lose
mulated in the day. During a typical fruit, whereas trees held at 21.1°C lost over
dark/cloudy/rainy day, adequate photosyn- 50% (Fig. 16.4; R.E. Byers, 2001, unpublished
thesis may occur to meet the carbon demands results). Thus, as light, darkness and temper-
during the day, regardless of the temperature ature change after the application, the tree
or the light level. However, the period during will become more or less susceptible to the
darkness may cause a critical demand for absorbed thinner.
photosynthates that are produced only during
the day. The hours from 0800 to 1700 h, there-
fore, must provide the excess photosynthates 16.7.2 Effect of post-application temperature
required for the night period (approximately on chemical fruit thinning
15 h each day from 1700 to 0800 h); thus, the
average night temperature, rather than the Olien and Bukovac (1978) found that warm
minimum temperature, may constitute a major temperatures were required (approximately
influence on fruit retention each 24 h period. 10°C, or an activation energy of 30–32 kcal
In several field experiments where trees mol1) for ethephon to break down and
were sprayed with carbaryl (or NAA) on the release ethylene in detached cherry leaves.
first day of an artificial shade period, trees lost These data suggest that applications of ethep-
more fruit than if trees were not shaded (Byers hon made under cool conditions (i.e. below
et al., 1971; Table 16.3). In addition, where arti- 10ºC) may not result in adequate ethylene evo-
ficially shaded trees were sprayed with car- lution to result in effective thinning. However,
baryl, less thinning occurred if 1 day of full when potted trees grown in the field that were
sun was allowed after the shade period but sprayed with ethephon were moved into
before carbaryl was applied. In these studies, controlled-environment rooms for 2 days, they
typically, 2 days of artificial shade induced were thinned equally well at 4.4 or at 21.1°C
more fruit drop than occurred following the (R.E. Byers, 2001, unpublished results).
application of the commonly used chemical Another group of potted trees did not give off
thinners (carbaryl or NAA) (Byers et al., 1991). ethylene from the ethephon spray when held
In three separate laboratory experiments, at 4.4°C for 5 days but, when moved to
fruit retention was inversely related to tem- another controlled-environment room at
perature when trees (with fruit of approxi- 21.1°C, ethylene was evolved at a level similar
Table 16.3. Effect of shade (92%) on chemical thinning of ‘Campbell Redchief Delicious’/MM.111 apple
trees (from Byers et al., 1991).
Shaded days (S) Fruit per cm2

Chemical thinners (T) cross-sectional
area of limb
Treatment1 34 35 36 37 38 39 40 (+ 56 days AFB)
Control (no shade) 7.93 a2

Carbaryl + oil T 4.34 b
Carbaryl + oil ST
ST SS 1.05 c
Carbaryl + oil S
S SS T 2.92 b
Carbaryl + oil S
S SS T 3.77 b
Shade S
S SS 4.42 b
Natural light levels 66 65 62 62 56 57 60
(PPF mol m2 day1)
1 Full
bloom occurred 22 April 1988. Fruit size was 22.4 mm ± 0.35 mm on 27 May 1988 (35 days AFB).
2Means of six trees and the control mean represented 767 fruit. Means separation within columns by
Duncan’s multiple range test, 5% level.
PPF, photosynthetic photon flux.
10 to those constantly held at 21.1°C. These data

4.4°C suggest that the response from ethephon thin-
9
Fruit per cm2 trunk cross-sectional area
a 21.1°C
ning sprays is largely controlled by the plant
8 metabolic status in response to changing envi-
ab
ronmental temperatures.
7 ab Personal observations of chemical thinning
6 abcd by growers have led me to believe that the
abc combination of chemical thinner, low light and
5 bcd warm night temperatures frequently causes
abcd
bcd
4 serious overthinning, whereas the combina-
tion of thinners applied under low light plus
3 bcd
cool conditions causes much less thinning and
2 an increase in the proportion of undersized
and pygmy fruit. In addition, natural ‘June
d
1 drop’ appears to be triggered by a low leaf-to-
0
fruit ratio in periods of low light plus warm
44 h 68 h 92 h 116 h 140 h temperatures (Byers et al., 1991), but, if tem-
Hours of darkness peratures are cool, little ‘June drop’ may occur.
These results also raise the question
Fig. 16.4. Influence of continuous 44 to 140 h of about the effectiveness of carbaryl, NAA
darkness, or 125 h of 21 h darkness + 3 h sunlight and 6-BA when temperatures are cold
each 24 h on fruit retention of ‘Golden (4.4°C). Theoretically, if a chemical thinner
Delicious’/M.27 trees held in controlled has been absorbed, it may be present in the
environment rooms at 4.4ºC or 21.1ºC. Trees held
tissue and a critical level of metabolism (dri-
in the dark from 68 to 140 h at 4.4ºC maintained
more fruit than those held at 21.1ºC. Trees held for
ven by temperature) may be required for the
68 or more h at 21.1ºC lost more fruit than trees plant to respond to absorbed chemicals.
held at 4.4ºC. Letters indicate mean separation by Cool temperatures may delay or interfere
Duncan’s multiple range test, 5% level. (From R.E. with action and increasing temperatures
Byers, 2001, unpublished results.) may promote abscission.
428 R.E. Byers
16.7.3 Physiological vs. temperature basis 16.7.4 Temperature probabilities for

for timing chemical applications thinning
Much emphasis has been placed on deter- An analysis of temperatures in Winchester,

mining the maximum effectiveness for each Virginia, from 1984 to 1993, showed that
chemical thinner based on the diameter of the during the 3-week period from 0 to 21 days
developing fruit. For NAA, the maximum AFB, the maximum temperatures for 3 days
sensitivity period for abscission has varied or more reached above 29.5°C in 7 out of 10
considerably in several reports while in oth- years. In the 1-week period from 15 to 21
ers there has been no apparent relationship days AFB (typically considered the optimum
over the period from petal fall (PF) to about thinning period at 8–12 mm fruit diameter),
15 mm fruit diameter (i.e. PF to about 25 days 3 or more days of temperatures above 29.5°C
AFB) (Donoho, 1968; Leuty, 1973; Marini, occurred in only 3 out of 10 years. To obtain
1996). Carbaryl also appeared to have a wide the same degree of probability for the 3-week
period (7 out of 10 years) as for the 1-week
range of effective thinning times, ranging
period, the temperature would have to be
from PF to about 20 mm fruit diameter (i.e.
dropped by 5.5°C to 24°C. Since the fruit
PF to 30 days AFB). Furthermore, ethephon
diameter, Julian date or days AFB were not
may cause fruit abscission from PF to 30 mm
found to be important factors (Byers et al.,
fruit diameter (i.e. PF to 40 days AFB) (R.E.
2000a), the data suggest that thinning should
Byers, 2001, unpublished results). Fluctuating be based on temperature considerations in
temperatures during the post-thinning the 3-week period, not fruit diameter or days
period have seldom been the focus of field AFB. In this analysis, the chemical combina-
experiments. Many of the early timing tion chosen for thinning was the most
studies could have been confounded by post- important factor, followed by temperature
temperature and sunlight levels, which were and, thirdly, the strain of ‘Delicious’.
seldom reported. Donoho (1968), who Unfortunately, light levels were not readily
reported light, temperature and humidity, available for the data for these experiments,
suggested two periods of maximum sensitiv- and a host of other influential factors may
ity for NAA. However, these trees may have have been significant, but none were pre-
been more susceptible to thinning since the sumed to be of primary importance.
two maximum-sensitivity periods also
correlated with two low-light periods.
Regression analysis of 20 years of thinning 16.7.5 Pruning and mechanical fruit thinning
trials on five spur ‘Delicious’ strains using 50
different spray treatments indicated that Dormant pruning reduces the number of
higher night temperatures were more highly flower and vegetative buds, resulting in a
correlated to chemical thinning (r2 = 0.19; greater supply of organic carbon reserves for
r2 = 0.21) than daytime temperatures (r2 = 0.12; fruit set and fruit growth for the remaining
r2 = 0.11) or the average of the day and night- vegetative and floral buds. Pruning may
time temperatures (r2 = 0.16) (Byers et al., easily remove 30–80% of the flower buds
2000a). Even though the regression coefficients before growth has started in the spring. In
were low, they were highly significant. In addition, pruning restricts tree height and
these data sets, chemical thinning was not sig- spread, reduces canopy density, increases
nificantly correlated with days AFB or fruit spray penetration, helps maintain tree struc-
diameter; however, a ranking of spray treat- ture, promotes regular bearing, stimulates
ments by various chemical combinations cho- shoot growth, inhibits flower-bud formation
sen for degree of expected thinning (r2 = 0.71) but increases spur vigour and flower-bud size,
was highly significant. In addition, a ranking increases the percentage of flowers setting
of the ‘Delicious’ strains by ease of thinning fruit, improves fruit size, quality and colour
was also highly significant (r2 = 0.15). and results in a reduction of the current
season’s yield but may promote yields of the for thinning apple and pear fruit were found
most valuable fruit sizes over the 2-year bear- to be non-selective for apple fruit size, and
ing cycle (Mika, 1986). Forsythe (1802, the trees must be pruned to have stiff
reviewed by Davis, 1957) made detailed rec- branches. Mechanical thinning has been
ommendations for pruning and training sys- found to be effective only for short, non-
tems designed to ‘keep trees in a constant state flexible peduncles, which are produced only
of bearing, which if left to nature would pro- on a few cultivars at about 60 days AFB. No
duce a crop only once in two or three years’. improvement in return bloom was found.
Roberts (1952) showed that fruit set, fruit size, Late hand-thinning or late chemical thinning
leaf size, shoot length and the subsequent sea- would be expected to increase fruit size due
son’s return bloom could be substantially to removal of smaller fruit.
increased by detailed removal of 70% of the
growing points by heading cuts in the dor-
mant season, even after flower-bud size had 16.8 Chemicals that Induce Flower-bud
been determined. He demonstrated that heavy Formation Without Thinning
pruning in the ‘on year’ could decrease the
number of flower buds and stimulate growth When biennial-bearing cultivars flower
and flowering in the ‘off year’. heavily and carry a full crop, early thinning
Mechanical methods of fruit (or flower) may not provide an adequate return bloom
removal have the advantage that the number for a return crop in the next season. During
and distribution of fruit remaining on the the thinning period, multiple applications at
tree can be visually observed during the low rates of ethephon have been used to pro-
thinning process. Mechanical shakers used mote return bloom of apple, while avoiding
1. Defruiting at bloom
2. Hand-thinning at bloom
3. Hand-thinning at 17.3 mm
4. Hand-thinning 50 days AFB
5. Ethephon + hand-thinning at 17.3 mm
6. Ethephon + CaNO3 + hand-thinning at 17.3 mm
14 7. Ethephon + 18N + 18P + 18K + hand-thinning at 17.3 mm
8. CaNO3 + hand-thinning at 17.3 mm
9. 18N + 18P + 18K + hand-thinning at 17.3 mm
10. Control + hand-thinning at 17.3 mm (60–70% FB in 1998) a
12
11. Control (0–10% FB in 1998)
Fruit cm–2 cross -sectional area
ab
10
(3 June 1999)
8
bc bc
6
cd
4
cd
2 d d
d
d d
0
1 2 3 4 5 6 7 8 9 10 11
Seven spray applications from PF to 49 mm in1998
(all treatments hand-thinned in1998)
Fig. 16.5. The combination of ethephon plus a nitrogenous foliar spray in the spring of 1998 (seven weekly
applications) provided adequate return bloom for a full crop in 1999 (treatment Nos 6, 7). ‘York’/M.27 trees
were selected for 90% or more of the spurs flowering. Trees with 60% spurs flowering or 10% spurs flowering
(treatment Nos 10, 11) also had an adequate return bloom for a full crop in 1999. Letters indicate mean
separation by Duncan’s multiple range test, 5% level. (From Byers et al., 2000b.)
430 R.E. Byers
fruit abscission from higher rates (Byers, able progress has been made to increase crop
1993). In one study when trees were heavily value by early flower and fruit thinning, con-
loaded, neither ethephon nor a foliar nutri- tinued efforts will be needed to develop as
ent spray alone promoted flower-bud forma- new cultivars, rootstocks and training sys-
tion, but the combination of the two greatly tems evolve.
promoted return bloom and fruit set (Byers To maximize crop value, future strategies
et al., 2000b; Fig. 16.5). There is some evi- may include: (i) pre-conditioning chemical
dence that NAA may directly promote spray(s) to make trees more responsive to
flower-bud formation (Harley et al., 1958) thinning chemicals; (ii) combinations of
and it has occasionally been used commer- chemicals with different modes of action
cially for that purpose. (photosynthetic inhibitors, auxin transport
inhibitors, flower bud inhibition or stimula-
tion, adjuvants, pruning and other cultural
16.9 Conclusion practices) that lead to consistent and reliable
cropping; (iii) accurate and continuous com-
The economic impact of thinning on crop puterized information on tree physiological
value is comprised of the yield and prices of condition (or carbon balance) and their
the most valuable fruit size categories aver- interactions with important environmental
aged over two or more years. Early fruit or fluctuations that influence natural fruit reten-
bloom thinning has not always achieved the tion/abscission and chemical response(s);
desired fruit size, return bloom, and/or and (iv) chemical application strategies to
annual bearing, particularly of strongly bien- removal of smaller unwanted or pygmy
nial bearing cultivars. Even though consider- fruit.
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17 Endogenous Hormones and Bioregulator

Use on Apples
Duane W. Greene
Department of Plant and Soil Sciences, University of Massachusetts, Amherst,
Massachusetts, USA

17.2 Classification and Chemistry of PBRs 438
17.2.1 Auxins 438
17.2.2 Gibberellins 439
17.2.3 Cytokinins 439
17.2.4 Ethylene 440
17.2.5 Abscisic acid 440
17.2.6 Aminoethoxyvinylglycine (AVG) (ReTain®) 441
17.2.7 Benzyladenine (BA) 441
17.2.8 2-Chloroethylphosphonic acid (ethephon, Ethrel®) 442
17.2.9 Indole-3-butyric acid (IBA) 442
17.2.10 Naphthaleneacetic acid (NAA) 443
17.2.11 Prohexadione-calcium (Apogee®) 443
17.3 Application of PBRs 443
17.4 Development and Maintenance of Tree Structure 444
17.4.1 Increased lateral branching 444
17.4.2 Suppression of water-sprout growth 445
17.4.3 Suppression of root suckers 445
17.5 Vegetative-growth Control 446
17.5.1 Growth control with Apogee® 446
17.5.2 Growth control with ethephon 448
17.6 Influence on Flowering and Fruit Set 449
17.6.1 Promotion of flowering in non-bearing trees 449
17.6.2 Promotion of flowering on bearing trees 449
17.6.3 Inhibition of flowering 449
17.6.4 Increasing fruit set 450
17.7 Control of Preharvest Drop 451
17.7.1 Control of preharvest drop with AVG 451
17.7.2 Control of preharvest drop with NAA 451
17.8 Improving Fruit Appearance and Shape 452
17.8.1 Improving fruit shape 452
17.8.2 Improving fruit finish 453
17.9 Influencing Fruit Maturity and Quality 454
17.9.1 Advancing fruit ripening with ethephon 454
17.9.2 Increasing red colour with ethephon 454
17.9.3 Delaying ripening and improving fruit quality with ReTain® 455

438 D.W. Greene
17.1 Introduction PBRs vary from country to country; some are

registered for use in limited regions of the
Nowhere in agriculture do plant bioregulants world. In this chapter, the main focus is on
(PBRs) play a more important role than in mainstream uses where their application is
apple production. While they are quite widespread and they represent an important
expensive, they are generally used on high- component in apple production.
value crops where the cost can be recovered This chapter, is organized by physiologi-
because of the higher value of the crop. cal responses that occur in the apple. Many
Frequently, PBRs do not increase yield or, if PBRs influence several systems in a plant.
they do, it is not the only benefit. The most Therefore, one compound may be mentioned
likely influence of PBRs is on improving fruit several times in this chapter, since it may
quality and increasing consumer appeal. have widespread effects on several different
PBRs, as discussed in this chapter, refer to physiological processes.
three types of compounds. First, there are the
hormones that occur naturally and are pro-
duced by the plant. Secondly, there is a 17.2 Classification and Chemistry of
group of synthetic plant growth regulators PBRs
that are not found naturally in any plants.
Thirdly, there are some compounds that are The growth of plants and the synchronous
structurally very similar to hormones but control of all physiological processes in the
they do not occur naturally in the target plant are directed and controlled by hor-
plants. Benzyladenine (BA) and indole-3- mones (Davies, 1995). These hormones are
butyric acid (IBA) are compounds that fit synthesized by the plant and, for different
into this latter category. PBRs, whether natu- classes or hormones, their chemical struc-
rally occurring or synthetic, generally act by tures are quite different. Hormone concen-
regulating or modifying natural physiologi- trations, activities and responses are
cal processes within the plant. influenced by the environment, cultural and
PBRs are not and should not be consid- management activities, pests and the appli-
ered to be pesticides, since they are not toxic cation of synthetic plant growth regulators.
nor do they act directly to affect either insects The purpose of the first part of this section is
or diseases. However, they can be considered to outline briefly classes of endogenous hor-
important components in an integrated food- mones in the plant and to indicate the physi-
production system, since they can influence ological processes where each play a key and
plant composition, which can have secondary pivotal role.
but important influences in controlling There are five generally recognized
insects and diseases. For example, growth classes of endogenous plant hormones. The
retardants can affect tree microclimate by chemical structure of the hormone or the pri-
allowing better light penetration and air mary example of the class is illustrated in
movement so that the fruit is less susceptible Fig. 17.1.
to fungal diseases, such as sooty blotch and
fly-speck. Furthermore, growth retardants
can make shoot tips less favourable for aphid 17.2.1 Auxins
infestation or infection from the bacterium
that causes fire blight (Yoder et al., 1999). The auxins were the first group of plant hor-
PBRs are used to control or influence mones to be discovered. Since there were no
many aspects of apple production, including protocols that had previously been estab-
control of preharvest drop, adjustment of lished and the very existence of compounds
crop load, regulation of flower-bud forma- that regulated plant growth had not been
tion, modification of tree structure, suppres- established, it took over half a century for
sion of growth, alteration of fruit shape, indole-3-acetic acid (IAA) to be discovered,
enhancement of fruit red colour and its role defined and its structure identified
improvement in cosmetic appeal. The uses of (Thimann, 1969). Although there are a num-
Endogenous Hormones and Bioregulator Use 439
CH3
H3C CH3 CH2 COOH
OH N
COOH H
O CH3
Abscisic acid Indoleacetic acid
CH3
O H N CH2 CH C
CH2OH
C O N
HO OH H N
COOH C CH2
H
Gibberellic acid (GA3) Zeatin
H H
C C
H H
Ethylene
Fig. 17.1. Structures of the major endogenous plant hormones.
ber of structurally similar compounds that centration are characteristic of that species
possess auxin activity, IAA is recognized as (Cleland, 1969). In apple, several gibberellins
the dominant auxin present in most plant have been identified, but gibberellin A4 (GA4)
species. It is produced in young leaves and and GA7 are generally present in the highest
in the shoot apex and moves only downward concentrations. Gibberellins are most noted
in the plant in the phloem. It plays an impor- for their ability to promote stem elongation,
tant role in a wide range of plant processes, an effect that can be very striking at times.
including apical dominance, fruit growth, Several commercial plant-growth retardants,
fruit set, root initiation, fruit ripening, leaf such as paclobutrazol, uniconizol, prohexa-
senescence and fruit and leaf abscission. dione-calcium and ancymidol, act by inhibit-
When auxins are exogenously applied, they ing gibberellin biosynthesis within the plant.
usually stimulate the production of ethylene; Gibberellins can inhibit flower-bud forma-
thus some auxin effects are manifested tion in many woody dicotyledonous species.
through ethylene responses. IAA is not used The inhibitory effect of fruit on return bloom
commercially but the synthetic auxins naph- in apple is attributed to gibberellins, pro-
thaleneacetic acid (NAA) and IBA are in duced in seeds, which move out and inhibit
wide commercial use. the formation of flowers for the following
season in the subtending bourse bud.
Gibberellins aid seed germination by stimu-
17.2.2 Gibberellins lating the conversion of starch to sugar.
This is a very large family with at least 121

different gibberellins having been identified 17.2.3 Cytokinins
in higher plants. Unlike the auxin IAA, which
is the dominant auxin in most plants, each The compounds in this group gained their
plant species has several gibberellins, and fre- generic name because they stimulate cell
quently the specific gibberellin and its con- division (cytokinesis) (Moore, 1989). The two
440 D.W. Greene
most common cytokinins are zeatin and its high rates, it can be a potent growth retar-
sugar conjugate, zeatin riboside. They stimu- dant. Commercially, ethylene is administered
late cell division in fruit, especially during to the plant as a spray solution of the plant-
the early stages of fruit development follow- growth retardant 2-chloroethylphosphonic
ing petal fall. Cytokinins are also used to acid (ethephon).
stimulate cell division and shoot generation
in tissue-culture propagation. Cytokinins
interact with auxins in the control of apical 17.2.5 Abscisic acid
dominance. They can overcome the
inhibitory effect of auxins on lateral bud Abscisic acid (ABA) was discovered inde-
development when applied exogenously or pendently in two laboratories in the 1960s
when produced endogenously in the roots (Moore, 1989). One laboratory identified it as
and translocated to the stems. Cytokinins a dormancy-inducing compound, while the
can delay or defer leaf senescence. They are other laboratory found it while studying
produced in all actively growing and divid- abscission in cotton. While it is associated
ing tissue, including leaves, shoot and root with dormancy, acts as a growth retardant
tips and seeds. However, many believe that and can cause abscission in selected crops, its
the roots are the most important site of syn- most important physiological function
thesis and, once produced, they are trans- appears to be in regulating water relations
ported upwards in the plant xylem. BA is a within the plant (Beyer et al., 1984). ABA is
synthetic cytokinin, and it is an active ingre- very closely associated with stomatal move-
dient in the commercial products Promalin® ment. When plants are stressed, ABA levels
and Accel®. increase and cations (especially potassium)
are pumped out of the guard cells, causing
stomata to close. Since plants have such an
17.2.4 Ethylene active and effective metabolic system for reg-
ulating ABA levels in the plant, exogenous
Ethylene is the only plant hormone that, in applications of ABA are so transient that
its natural state, is a gas. For many years, ABA is not used commercially to regulate
plant physiologists refused to acknowledge plant water relations.
the hormonal status of ethylene, because it The second and largest group of PBRs
was a gas (Abeles, 1973). Ethylene moves that are in general use in agriculture are syn-
easily within the plant through intercellular thetic organic compounds. The chemical
spaces and in a dissolved form in the cyto- structures and the trade name, chemical
plasm (since it is quite soluble in water). name and manufacturers of some important
Frequently ethylene is stored within the PBRs used on apples are illustrated in Fig.
plant as its precursor 1-amino-cyclo- 17.2 and Table 17.1, respectively. They fall
propane-1-carboxylic acid (ACC). Conver- into several categories. Some very closely
sion from ACC to ethylene occurs following resemble structurally endogenous hor-
an environmental stress or triggering by an mones. They are generally more effective
internal physiological signal. Ethylene plays than the endogenous compound they resem-
a key role in fruit ripening and abscission. ble, in large part because the regulatory sys-
Apple is a climacteric fruit and it generates tem within a plant is not nearly as efficient
a large amount of ethylene as it enters the in breaking down, metabolizing or inactivat-
ethylene climacteric phase at maturity. The ing these compounds with their slightly
ethylene given off in this process stimulates altered structure. A second category of com-
ripening and, when translocated to the pounds includes those that inhibit the
abscission zone in the pedicel, it initiates biosynthesis or transport of endogenous
biochemical changes that result in the hormones. These are also structurally unre-
destruction of cells in the abscission zone. lated compounds that directly or indirectly
Ethylene promotes flower and fruit senes- affect change within the plant and alter
cence and induction of flowering and, at physiological responses.
O
CH2COOH
C O
HO OH
COOH C CH2
Gibberellin A4 Naphthaleneacetic acid
O HN CH2
N N
Cl CH2 CH2 P O–
O– N N
2-Chloroethylphosphonic acid 6-Benzyladenine
O– O
O NH3+
Cl–
O Ca++
O
+
O–
NH3 –O
O
Aminoethoxyvinylglycine Prohexadione-calcium
Fig. 17.2. Structures of the most commonly used plant bioregulants (PBRs) on apples.
17.2.6 Aminoethoxyvinylglycine (AVG) ReTain® in 1997 for preharvest drop control

(ReTain®) and improvement of fruit quality in apples.
AVG is a naturally occurring amino acid that

was first discovered by scientists at Hoffman 17.2.7 Benzyladenine (BA)
LaRoche in the early 1970s. Its primary mode
of action is to inhibit ethylene biosynthesis The ability of BA to increase the length of
(Boller et al., 1979). It blocks the enzyme ACC apples was recognized by Williams and Stahly
synthase, a key enzyme in the ethylene (1969) several decades ago. Scientists at Abbott
biosynthetic pathway. AVG elicits many Laboratory combined it with GA4+7 in equal
responses in apple trees, including increas- amounts in the proprietary product Promalin®
ing fruit set, increasing vegetative growth, specifically to elongate ‘Delicious’ apples and
increasing fruit length/diameter (L/D) ratio, to promote branching on apple trees. When BA
retarding fruit ripening, retarding preharvest was evaluated independently, it was found that
drop and stimulating branching. Efforts to BA had several properties that are characteris-
register this compound in the early 1980s tic of a good chemical thinner, including abil-
were not pursued because of economic rea- ity to promote abscission, increase flower-bud
sons. Following the loss of daminozide as a formation, increase fruit size independently
preharvest fruit-drop control compound, of thinning effects and increase fruit firmness
Abbott Laboratories resumed development in some instances. An altered Promalin® for-
of this compound for commercial use. AVG mulation containing primarily BA was regis-
received full label registration as the product tered as Accel® for use on apples in 1994.
442 D.W. Greene
Table 17.1. Common plant bioregulants (PBRs) used in apple production.
PBR Common/trade name Chemical name Manufacturer/seller
AVG Retain® Aminoethoxyvinylglycine Valent USA Corporation

BA Cylex™ N-(phenylmethyl)-H-purine-6-amine Valent USA Corporation
GA4+7 Provide® Gibberellins A4+7 Valent USA Corporation;
Regulex® Valent USA Corporation;
TypRus™ Nufarm Americas Inc
GA4+7 + BA Promalin® Gibberellins A4+7 + N-(phenylmethyl)- Valent USA Corporation;
1.8% + 1.8% Cytolin® H-purine-6-amine Valent USA Corporation;
Pomina® Valent USA Corporation;
Typy™ Nufarm Americas Inc;
Perlan® Fine Agrochemicals Ltd
BA + GA4+7 Accel® Gibberellins A4+7 + N-(phenylmethyl)- Valent USA Corporation;
1.8% + 0.18% RiteSize™ H-purine-6-amine Nufarm Americas Inc;
Ethephon Ethrel® 2-Chloroethylphosphonic acid Adventis Crop Protection;
Ethephon 2 Micro Flo Company LLC
NAA Fruitone® N Naphthaleneacetic acid Amvac Chemical Corporation
K-Salt™ Fruit Fix™ 200 Naphthaleneacetic acid potassium
salt
K-Salt™ Fruit Fix™ 800 Naphthaleneacetic acid potassium
salt
Tree-Hold A-112 Naphthaleneacetic acid ethyl ester
NAAm, NAO Amid-thin® Naphthaleneacetamide Amvac Chemical Corporation
Prohexadione- Apogee® Calcium 3-oxido-5-oxo-4-propionyl- BASF Corporation
calcium Regalis® cyclohex-3-ene-carboxylate
17.2.8 2-Chloroethylphosphonic acid ily absorbed and readily moves into the cyto-
(ethephon, Ethrel®) plasm, where the pH level is slightly below
neutrality (pH 7). Ethephon is unstable at this
Ethephon was the first ethylene-based plant- pH range and autocatalytically breaks down
growth regulator to be available in the mar- to liberate ethylene gas within the cell. The
ket. It was introduced in 1971 to stimulate ethylene liberated from the breakdown of
latex flow in rubber trees. Since that time, its ethephon frequently stimulates the plant to
use has been considerably broadened and it produce even more endogenous ethylene.
has become one of the most useful and
diversified PBRs applied. It is now registered
on over 20 horticultural and agronomic 17.2.9 Indole-3-butyric acid (IBA)
crops, including fruit crops, such as apple,
cherry, pineapple and grapes. Soon after the discovery of auxins, the root-
Ethylene is a hormone that influences sev- promoting activity of IBA was recognized. It
eral physiological systems within a plant. has been used for many years to stimulate
However, the problems associated with rooting in tissue culture and in shoot cut-
administering ethylene in its gaseous form are tings taken for vegetative propagation
immense. Ethephon simplifies this and makes (Hartmann and Kester, 1983). It remains
it extremely easy and convenient to apply pre- today as the only hormone, or as a major
cise doses of ethylene when needed. Ethephon component in several commercially available
is applied as an aqueous spray, where it is eas- rooting formulations, for root induction.
17.2.10 Naphthaleneacetic acid (NAA) 17.3 Application of PBRs
NAA was one of several auxins that were The majority of PBRs are applied as a foliar
identified as retarding preharvest drop in spray and this has historically been done
apples prior to harvest, while also promot- with a dilute application, where the spray
ing abscission on young developing fruit material is applied to near the drip (or
when applied soon after bloom. It is used runoff) point. This was an effective approach
to promote rooting in some rooting formu- to ensure that an appropriate distribution of
lations. It is the only auxin to have sur- the active ingredients are uniformly distrib-
vived over 50 years of regulatory scrutiny uted within and to the tops of large trees
and remain registered for use today both propagated on seedling or vigorous clonal
as a chemical thinner and as an inhibitor of rootstocks. The trend for the past two
preharvest drop on apples. decades has been to grow apple trees as
smaller-stature trees at much higher densi-
ties. The use of larger equipment designed to
17.2.11 Prohexadione-calcium (Apogee®) deliver spray to the tops of large trees is
inappropriate in high-density plantings
Several commercially-produced PBRs are because of the large amount of space
available that interfere with gibberellin required between rows. Accompanying
biosynthesis. However, Apogee® represents increased tree density has been a reduction
a new class of gibberellin inhibitor, which in the size of equipment used and a reduc-
inhibits GA biosynthesis at a later stage in tion in the volume of spray used to deliver
the biosynthetic pathway and thus offers both pesticides and PBRs.
the potential of acting with fewer side- Byers et al. (1971) first introduced the con-
effects (Rademacher, 1991). Apogee® cept of tree-row volume (TRV) to determine
retards vegetative growth on apples, which the volume of spray necessary to achieve
improves spray penetration and coverage, adequate coverage of trees planted at higher
reduces pruning time and provides a more densities. An example of a dilute TRV calcu-
open canopy that facilitates light penetra- lation for an apple orchard is illustrated in
tion into the tree. Table 17.2. The concept has evolved and been
Table 17.2. Calculations of dilute volume necessary to wet apple foliage in an orchard to saturation.
Parameters to measure
Canopy height (m) – distance from first scaffold to the top of the canopy
Tree width (m) – average maximum width of a tree from branch tip to branch tip
Area of a hectare (m)
Row length = –––––––––––––––––
Row width (m)
Tree-row volume (TRV) or canopy (m3) =
Canopy height Tree width Row length
Dilute volume requirement per hectare equals about 1 l of spray for each 10 m3 of foliage
Example calculation of dilute spray requirement
Trees in a sample orchard have a canopy height of 3.75 m, a tree width of 4 m and a row width of 6 m.
10,000 (m2 ha1)
TRV = 3.75 4 ––––––––––––– = 25,000
6
TRV
Dilute volume = ––– = 2500 l ha1
10
444 D.W. Greene
refined in recent years. Effective and consis- spray deposition in the tops and interior of
tent use of PBRs must take TRV into account trees is often less than satisfactory.
if spray applications of PBRs are applied at
reduced spray volumes (Bukovac, 1980). For
example, label recommendations for the use 17.4 Development and Maintenance of
of AVG as a preharvest-drop-control com- Tree Structure
pound are given in grams of active ingredient
per hectare. However, this bioregulant may The fruiting structure of an efficient apple
be applied on blocks that have a TRV ranging tree is developed during the formative years.
from 935 to 3800 l ha1 (100–400 gal acre1). This fruiting structure should have several
Since the drop response to ReTain® is linear characteristics: it should be developed
with the dose sprayed on the tree, one can rapidly, make efficient and effective use of
expect a much stronger drop-control the allotted space, efficiently intercept the
response on trees with the lower TRV. In this available sunlight and possess a structure
case, four times as much material is being that encourages the development of an
applied on the leaves and fruit of smaller appropriate balance between fruiting and
trees than on the larger ones. When applying vegetative growth (Forshey et al., 1992).
PBRs at less than dilute TRV, it is always pru-
dent to first calculate TRV and then put in the
spray tank the amount of material suggested 17.4.1 Increased lateral branching
on the label for the area being sprayed. The
surfactants and other spray additives to aid Many cultivars, especially those that are tip
penetration are generally not concentrated. bearers or those that possess a spur-type
There are several problems associated with growth habit, do not naturally form an ideal
the application of PBRs in low-volume sprays structure or have an appropriate growth dis-
and these may result in variable responses tribution (Elfving, 1984; Miller and Eldridge,
(Bukovac, 1984, 1985). First, the chance of 1986). Many cultural and management tech-
over- or under-application is accentuated since niques are currently in use or have been eval-
the response to most PBRs is linear with con- uated to improve tree structure. These include
centration or dose. When PBRs are applied as pruning, bending, spreading, notching,
a dilute spray, the chance of over-application deblossoming and bud removal. In general,
is minimized because excess spray will drip branching in young trees is inhibited because
off the tree. Even if an error of 10% in the the apical buds on a tree or shoot suppress
amount of PBR applied is made, consequences bud break and shoot growth of buds in lower,
are generally not great. However, if a spray is more basal positions. Complete removal of
applied in one-sixth to one-tenth the volume the shoot tip by pruning usually results in the
of water used for a dilute spray (six- to ten- growth of two or three undesirable shoots
fold) and the same 10% error is made, this with sharp branch angles immediately behind
error can be magnified 600–1000%, with the cut (Forshey et al., 1992). Alternatively,
resulting consequences of over- or under- Promalin®, a proprietary product containing
application. Secondly, where a lower volume equal amounts of GA4+7 and BA, may be
of water is used, the drying time of the spray applied to apple trees shortly after bloom to
will generally be shorter. It is generally encourage lateral bud growth and to improve
accepted that there is reduced uptake of foliar- tree structure (Table 17.3). This product may
applied substances under such circumstances. be used on trees where the apical bud has not
Therefore, efficacy, even when the same been removed by pruning. Consequently, the
amount of PBR is applied, may be diminished undesirable regrowth following pruning can
due to reduced uptake. Thirdly, smaller be avoided. It is most successful when used
droplet sizes are generally associated with on healthy, rapidly growing trees. It generally
methods that apply lower spray volumes. requires application of from 125 to 500 mg l1
Smaller droplets generally do not carry as far with an effective surfactant, made when ter-
and are more affected by wind. Therefore, minal growth is between 25 and 75 mm in
Table 17.3. Influence of BA + GA4+7 and BA alone (application when shoots of ‘Macspur McIntosh’/M.26
were 3 cm in length) on lateral branching and shoot growth (from D.W. Greene, 1982, unpublished results).
Spurs Lateral Mean lateral Blossom clusters

Treatment per shoots per shoot length per cm LCSA
(mg l1) cm LCSA cm LCSA (cm) year after application
Control (0) 4.1a 0.7b 24.0a 11.9a

GA4+7 + BA 500 2.1b 2.6a 20.6b 0.1b
BA 500 2.7b 2.3a 17.0c 13.8a
Values with uncommon letters are significantly different at odds of 19 to 1.

LCSA, limb cross-sectional area.
length (Forshey, 1982). Rates that effectively opment of insects and diseases. They can be
stimulate lateral branching are usually high removed by hand, but at considerable
enough to completely defruit trees and the expense. They can be pruned out but regrowth
presence of gibberellins in this product is virtually ensured when using this approach.
inhibits flowering for the following year Control of the growth and development
(Table 17.3). Therefore, the use of Promalin® is of water sprouts is most easily accomplished
limited to situations where fruiting is neither with NAA. The Tree-Hold Sprout Inhibitor®
anticipated nor desired for the year of appli- (Amvac, Los Angeles, California) is specially
cation and the year after application. formulated, using the ethyl ester of NAA for
Branch development can be a somewhat sucker control, and it is more effective than
random event. Generally, buds that grow other formulations of NAA, such as the
into good lateral shoots come from large sodium salt, which is used for chemical thin-
buds and they are located either laterally or ning (Raese, 1975). Generally, this product is
on the top of a branch. The chance of stimu- mixed with 25–50% interior white latex
lating branching on a limb portion can be paint, to give a final NAA concentration of
improved immensely by notching buds prior 0.5–l.0% (Miller and Ware, 1980). The latex
to Promalin® application. Notching is done paint marks and identifies the area treated
using a hacksaw blade to remove a strip of and thickens the mixture so that it remains
bark one-third around the stem immediately on the treated area. It may be applied with a
above the bud that one wishes to develop brush, paint roller or hand-pump sprayer
into a lateral shoot (Greene and Miller, 1988; with a sponge attached to the nozzle.
Greene and Autio, 1994). This can be done a The NAA should be applied after prun-
few days either before or after full bloom. ing, during the dormant period. Both the cut
surfaces and 50–75 mm surrounding the cut
should be treated. However, whole branches
17.4.2 Suppression of water-sprout growth should not be treated. As a useful guide, no
more than 10% of the total limb surface area
As apple trees mature, it is frequently neces- should be treated. Application should be
sary to make pruning cuts to restrict tree made no later than bud break, because of the
height or to contain trees within their allotted possibility of thinning, due to NAA volatil-
space. These cuts often stimulate the develop- ity, and reduced fruit size, due to a direct
ment and rapid growth of upright shoots that effect of NAA.
originate from latent buds within the wood
(Forshey et al., 1992). They are referred to as
either water sprouts or suckers. They are very 17.4.3 Suppression of root suckers
undesirable because they are unproductive,
create unwanted shade within the canopy, Many of the most popular dwarfing root-
impede distribution and coverage of spray stocks and interstem trees develop suckers at
materials and can foster the growth and devel- the base of trees. These unsightly shoots
446 D.W. Greene
serve as an entry way for fire blight into restricted fertilization and the control of
trees, harbour insects and diseases, make water (Ferree, 1981). Used alone, many of
some orchard activities such as mowing dif- these techniques may be only marginally
ficult and, if unattended to, will grow up effective. The two PBRs that hold the greatest
into a tree. Control of these is accomplished commercial potential to control vegetative
either with the use of NAA ethyl ester as the growth on apples are Apogee® and ethephon.
Tree-Hold Sprout Inhibitor® formulation
(Miller, 1977) or in Europe and the southern
hemisphere with Tipoff® (Midox Ltd, Kent, 17.5.1 Growth control with Apogee®
UK), which contains the free acid of NAA in
an emulsion with decanol. Root suckers are Apogee® is the newest growth retardant to
first cut to the ground, generally during the receive full label registration. Like several
dormant season and when the new root other growth-retarding chemicals, it acts by
suckers are 10–30 cm in height, they are inhibiting gibberellin biosynthesis (Evans et
treated (Miller, 1989). It is recommended that al., 1999). However, it is different in that it
the period from bloom to 4 weeks after blocks the gibberellin biosynthetic pathway
bloom should be avoided, because the at a point different from other known GA
volatility of NAA may cause fruit thinning. inhibitors. Once applied, it requires between
Tree-Hold® is prepared as described for con- 10 and 14 days to slow growth. It degrades
trol of water sprouts, except that water is within the tree in a few weeks, so repeat
used instead of the latex paint, while when applications are usually necessary to main-
Tipoff® is used it is diluted with two parts tain growth control throughout the whole
water. Both compounds are sprayed on the growing season (Byers and Yoder, 1999).
foliage of the root suckers. This should only Patterns of terminal growth and fruit-set
be done under calm wind conditions and characteristics differ among fruit-growing
using a sprayer that does not generate many regions, and the response to Apogee®
small droplets that will drift. Since thorough appears to differ depending upon the area of
coverage is critical and weeds may protect the country where it is used (Unrath, 1999).
root suckers from the spray, application of a Therefore, regional interpretation of the label
contact herbicide, such as gramoxone, a few is necessary to get the maximum response
days before application may improve con- desired.
trol. The initial application of Apogee® is
made as soon as there is sufficient leaf area
to absorb the chemical – generally at late
17.5 Vegetative-growth Control bloom or petal fall, when shoots are
25–75 mm in length. The growth response of
There is a very delicate balance between veg- ‘Macoun’ apple trees to one petal-fall appli-
etative growth and cropping in apple trees. cation of prohexadione-calcium at a high rate
This balance may be disrupted by abnormal or three applications at a lower rate is illus-
or catastrophic weather or by management trated in Fig. 17.3. It has no detrimental
mistakes that reduce fruit set or crop load effects on bees so the first application can be
and result in excessive vegetative growth. made even before bees are removed from the
Vigorous growth can negatively influence orchard. The initial application should be at
fruit quality, productivity, pest control and a TRV-adjusted concentration of 82–125 l1.
profit. Failure to control vegetative growth Application can be concentrated as long as
adequately in high-density blocks may ulti- coverage is uniform. Apogee® should be
mately result in the necessity to remove the applied with a surfactant at a rate of
block because of poor fruit quality and low 0.5–1.0 ml l1 to assure wetting and good
productivity. There are a number of no-chem- coverage. Apogee® will precipitate out in the
ical ways to restrict vegetative growth, spray tank if the water it is applied in is
including dormant, summer and root prun- ‘hard’ (i.e. has a high pH and contains high
ing, ringing and scoring, limb spreading, levels of calcium carbonate). Under these
40
35
Terminal growth (cm)

30
25
20
15
10
Control
5 Prohexadione-Ca 270 mg l–1
Prohexadione-Ca 90 mg l–1
0
0 20 40 60 80 100 120 140 160
Days after application
Fig. 17.3. Growth control of ‘Macoun’/M.7 apple trees using one petal-fall application of prohexadione-Ca
at 270 mg l1 or three applications of 90 mg l1 at approximately 2-week intervals starting at petal fall (after
Greene, 1999).
circumstances, the label recommendation is However, a consensus is emerging where

to add an equal weight of ammonium sul- the scenario that will probably be adopted
phate in the spray tank with Apogee®. The by most growers is to make an initial appli-
prohexadione-calcium formulation being cation at 25–50 mm of growth using 82–125
prepared for sale in Europe, and perhaps mg l 1. If a block is vigorous and the grower
other countries, contains ammonium sul- does not want to assume the responsibility of
phate; thus additional ammonium sulphate monitoring the resumption of growth on a
will not be necessary. nearly daily basis, a second application of
Prohexadione-calcium may increase fruit 62–82 mg l1 should be made 2 weeks after
set. This response is particularly pronounced the first. That may be all of the prohexa-
in areas that experience cool weather during dione-calcium required in cooler northern
the post-bloom period. Its effect is linear regions where terminal growth stops early
with increasing concentration (Greene, 1999). (Greene, 1999). Additional applications of
In the majority of cases where 250 mg l1 is 62.5–125 mg l1 may be required in warmer
used, fruit set will be increased, with a corre- growing areas with a longer growing season
sponding decrease in fruit size. In these (Unrath, 1999).
cases, more aggressive chemical thinning is
necessary to reduce crop load down to
17.5.1.1 Use of prohexadione-calcium to
desired levels.
control fire blight
Growers are given a number of treatment
choices involving concentration and num- Prohexadione-calcium will control fire blight
bers of applications to make under several on shoots by inducing resistance in the tree
growth-control circumstances. Experts who (Yoder et al., 1999). For it to be effective, it
have worked with this compound for several must be applied and growth retardation
years do not agree on the amount to apply must occur before infection (Table 17.4).
initially, how many applications to make and Generally, this requires that application
how to decide when it is appropriate to must occur a minimum of 10–12 days before
make additional applications. Growers may infection. The active ingredient in prohexa-
be required to use the trial-and-error method dione-calcium appears not to have any
initially to determine the best combinations direct effect on the fire blight bacteria. It is
for use in their situation. not effective on blossom blight, so tradi-
448 D.W. Greene
Table 17.4. Suppression of fire blight incidence and canker length by applications of prohexadione-Ca
and streptomycin to ‘Golden Delicious’ apple trees, Winchester, Virginia, 1997 (from Yoder et al., 1999).
% Inoculated shoots
Rate (mg l1) and timing infected 17 July; Mean canker
inoculated on: length (cm)
Prohexadione-Ca Streptomycin 17 July of shoots
21 May 27 May 28 May 4 June inoculated 4 June
0 0 48a 75d 12.1b

125 0 32ab 20b 2.2a
250 0 25ab 13b 0.4a
0 100 18a 50c 4.8a
250 100 16a 2a 0.7a
tional measures using streptomycin are Occasionally, the fruit crop on blocks of
appropriate. Application of prohexadione- bearing trees is lost due to frost. There is a
calcium to control fire blight should be delicate balance between vegetative growth
made at the same time as applications to and fruiting, and the high vigour caused by
control growth, when shoots are 25–50 mm crop loss may tip the balance so far in favour
in length, at a rate of 125 mg l1. A second of vegetative growth as to make it difficult to
application may be required 3 weeks later to bring the planting back into an appropriate
get the best results. balance between vegetative growth and fruit-
ing in successive years. Ethephon applied at
500 mg l1 when terminal shoots are
17.5.2 Growth control with ethephon 10–15 cm in length may be useful in retarding
growth and preserving an appropriate bal-
Ethephon is an extremely effective growth ance between vegetative growth and fruiting
retardant. It is also an effective fruit thinner, in the year that frost damage occurs. A sec-
and it can thin over the longest span of ond application of ethephon later, at a lower
developmental stages of any of the commer- rate, may be appropriate in warmer areas
cially used chemical thinners (see also with a longer growing season (Greene,
Chapter 16). Because it acts both as a growth 1996b). Growth retardation and increased
retardant and as a thinning agent, care must return bloom following ethephon application
be exercised to minimize thinning effects and compared with scoring on ‘Delicious’
when used on bearing trees. apple trees are illustrated in Table 17.5.
Table 17.5. Influence of ethephon application or scoring 12 days after

bloom on terminal growth and flower-bud formation the year after
application on 6-year-old ‘Richared Delicious’/M.7 (from Greene and
Lord, 1978).
Blossom clusters
Treatment Terminal growth per cm LCSA
(mg l1) (cm) year after application
Control 55a 3.0c

Ethephon 500 38b 7.2b
Ethephon 1000 35b 9.9a
Scoring 38b 8.8ab

17.5.2.1 Postbloom application on non- Often this is all the intervention necessary to
bearing trees ensure good repeat bloom. However, there
are situations where further promotion of
On non-bearing trees ethephon should be
flowering is appropriate. Growers have two
applied at 250–500 mg l1, when shoot
PBRs available to promote flowering on
growth is 7–12 cm in length. Growth control
bearing trees.
is more difficult on non-spur cultivars and
on trees budded on vigorous rootstocks. NAA may be applied after the time fruit
Increased flower-bud formation can be are susceptible to thinning but before the
expected the following year. flower-bud initiation period has ended,
Ethephon may be used on bearing trees using 3–5 mg NAA l1. The exact time will
but precautions must be taken to minimize vary from year to year but this period would
thinning if a reduction in crop load is not roughly range from the time fruit are 20 mm
desired. One strategy to avoid thinning is to in diameter to 8 weeks after bloom. The use
wait until the end of June drop (northern of NAA at these low rates is unlikely to affect
hemisphere) before applying ethephon. fruit size.
Using this approach, growth control will be Multiple doses of ethephon at 100–200 mg
less than if applied earlier, little or no thin- l1 may also be applied, starting at the end of
ning will take place and some increase in June drop. Byers (1993) enhanced flowering
return bloom will occur. These treatments on ‘Starkrimson Delicious’ by applying
may, however, advance ripening, which may either 12 weekly or six biweekly sprays of
result in early preharvest drop and early either 100 or 200 mg ethephon l1, respec-
breakdown of starch within the fruit. tively. Some growth control was achieved
with these treatments and fruit ripening was
advanced, as determined by starch rating.
17.6 Influence on Flowering and Fruit Set
17.6.1 Promotion of flowering in non- 17.6.3 Inhibition of flowering

bearing trees
Regulation of flowering and the maintenance
Ethephon is registered to promote flower-bud of an appropriate balance between vegeta-
formation on young bearing trees when tive growth and flowering is usually accom-
applied at 300–600 mg l1 shortly after bloom. plished on bearing trees by chemical
However, the apple trees now being planted thinning during the first 3 weeks after bloom
are being established at higher densities and on heavily blooming trees. Young develop-
are being propagated on dwarfing rootstocks. ing fruit are removed during this time, leav-
They generally bloom and set fruit early, ing some spurs with no fruit, which will
reducing the need to promote further flower- allow flower-bud formation for the following
ing. Since early production is a key to success year. It is possible similarly to establish an
in these blocks, the use of flower-bud-promot- appropriate balance between vegetative
ing sprays is usually not necessary. The use of growth and fruiting by inhibiting flower-bud
ethephon to promote flowering on non- formation during times when there is
bearing trees is generally restricted to vigor- reduced flowering on bearing trees.
ous rootstock/scion combinations, where Gibberellins are produced by the seeds.
flowering can be delayed for several years. These migrate out and move to the bourse
bud, where they inhibit flowering for the fol-
lowing year. Commercially available formu-
17.6.2 Promotion of flowering on bearing lations of GA3 (ProGib®) and GA4+7
trees (Provide®) may be applied during the first 4
weeks after bloom, at rates of 50–200 mg l1,
Flowering on bearing trees is most often reg- and this inhibits flower-bud formation for
ulated by promoting fruit abscission through the following year (Greene, 1989). This has
the use of chemical thinners (Chapter 16). not become a viable way to regulate flower-
450 D.W. Greene
ing in bearing trees because of several nega- be due to nutritional or physiological status
tive side-effects. GAs reduce seed number that result in stress. PBRs in these circum-
and seeds are important to aid the uptake stances may be used to enhance fruit set.
and mobility of calcium to the fruit within Mixtures of chemicals containing auxins,
the tree (Bamlage et al., 1990). Inhibition of gibberellins and cytokinins have been used
flowering by the use of GAs has not been in Europe to increase fruit set on pear trees
sufficiently perfected for precise and pre- that have been frost-damaged, but the
dictable reductions in flowering to be response is less predictable on apples
achieved (Greene, 2000). Poor pollinating (Goldwin, 1986). Gibberellins may increase
weather may occur at bloom, which would fruit set on apples but their application is
make it advantageous to have more, rather unlikely to become a widespread commercial
than less, bloom to get a commercial crop. practice for several reasons (Dennis, 1973).
Inhibition of flowering may be a viable The fruit-set response is erratic and unde-
technique on young trees that are just starting pendable, and the fruit that are set as a result
to come into production. The majority of of GA application are generally small and
apple trees being propagated in the USA and have a reduced storage potential. Further-
elsewhere are propagated on very precocious more, GAs inhibit flower-bud formation for
rootstocks. Early flowering and fruit set can the following year.
slow tree growth to the extent that trees are Prohexadione-calcium, when applied to
prevented from filling their allotted space control terminal growth and retard fire
and never reach their full yield potential. blight, may increase fruit set (Greene, 1999).
Furthermore, heavy set on young trees may The response is most prominent when rates
cause leader or limb breakage, which can above 125 mg l1 are used, when it is used in
result in substantial structural damage to the cooler climates or when cooler weather con-
tree. Sprays of 250–500 mg GA4+7 or GA3 l1 ditions occur during the post-bloom period.
can substantially reduce return bloom on Increased fruit set following Apogee® appli-
young apple trees, thus improving tree growth cation usually results in reduced return
(Table 17.6; Unrath and Whitworth, 1991). bloom the following year.
It is well known that AVG can increase
fruit set on apple (Williams, 1980). Generally
17.6.4 Increasing fruit set rates between 125 and 500 mg l1 are
required. Application can be made shortly
Even though trees may flower, there are before harvest or soon after bloom but before
occasions when these flowers fail to set and the start of June drop. It can increase fruit set
develop into a commercial crop. This may on young apple trees that are just starting to
occur because of a weather event such as a come into production but, as with other ways
frost or poor pollinating weather, or it may of increasing set, it frequently reduces return
Table 17.6. Influence of GA4+7 application made 4.5, 9, 13 and 18

weeks after petal fall on return bloom of ‘Redchief Delicious’/M.7 apple
trees (from Unrath and Whitworth, 1999).
Gibberellin Blossom clusters Bloom

concentration per cm LCSA suppression
(mg l1) the year after application (%)
0 7.8a 0
250 0.4b 95
500 0.1c 99
Values with uncommon letters are significantly different at odds of 19

to 1.
bloom for the following year. Increased fruit action of AVG is to inhibit ethylene biosyn-
set with the currently available commercial thesis in the fruit. In the fruit treated with
formulation of AVG is not on the ReTain® AVG, ethylene production will be reduced,
label at this time and, unless the cost of this synthesis of enzymes responsible for the
product is reduced, increased fruit set is not destruction of cells in the abscission zone
likely to be a use of this product. will be delayed and the fruit will remain on
The growth retardant paclobutrazol the tree for a longer period of time.
(Cultar®) when applied as a foliar spray can Label recommendations for the use of
increase fruit set the year following applica- AVG suggest that it should be applied at a
tion (Miller and Sweitlik, 1986). Increased rate of 125 g (a.i.) ha1, regardless of tree size
fruit set can occur for several more years if or TRV. However, the effectiveness of AVG as
higher rates are used or application is made a drop-control compound is linear with the
as a soil drench. Extreme care must be taken amount applied. Therefore, better drop con-
when using this compound, since residues trol can be expected on smaller trees in
may remain in the ground and retard growth blocks with a low TRV, simply because more
subsequently for several years. compound is delivered per area of tree than
on larger trees.
The suggested time of application of AVG
17.7 Control of Preharvest Drop is 4 weeks before anticipated normal harvest.
Applications made earlier than this run the
One of the greatest problems that apple risk of losing drop control before harvest
growers face is the drop of fruit before it can and, if application is delayed, fruit effects
be harvested. Some cultivars, such as will be diminished, some drop may occur
‘McIntosh’, are particularly prone to this before drop control is effective and harvest
problem. Drop on many cultivars can exceed may be unnecessarily delayed, because a
20%, but on the drop-prone cultivars it is not preharvest interval of 28 days is required
uncommon to have losses exceed 50% between application and harvest. The sili-
(Greene, 1996a). In areas where one or two cone-based surfactants ‘Silwet’ and ‘Sylgard’
cultivars are dominant, growers are fre- should be included at 0.05–0.1% to improve
quently faced with the challenge of harvest- the effectiveness. Concentrate application
ing a large portion of their crop in a short can be done as long as the spray volume is
period of time before fruit condition declines large enough to ensure both good coverage
and fruit are lost due to drop. Warm weather and adequate spray distribution within the
before harvest, drought stress and foliage tree. The effectiveness of AVG and NAA as
damage from insects and diseases all drop-control compounds on ‘McIntosh’
increase drop (Byers, 1997). Several com- apples is illustrated in Fig. 17.4. Rain soon
pounds have been used over the past half- after application may reduce the response to
century to control preharvest drop, but, due AVG. However, if one of the recommended
to regulatory reasons, only AVG (ReTain®) silicone-based surfactants is used and AVG
and NAA are currently available for wide- completely dries on the foliage, rain will
spread commercial use. have little effect, if any, on reducing prehar-
vest drop (Greene et al., 2000).
17.7.1 Control of preharvest drop with AVG

17.7.2 Control of preharvest drop with NAA
As apples ripen, they produce ethylene in
large amounts, which moves through the It has been known for over a half a century
intercellular spaces in the fruit and through that auxins can inhibit abscission. Many syn-
the pedicel to the abscission zone, where it thetic auxins inhibit preharvest drop and
stimulates the synthesis of enzymes, which several of these have been used commer-
ultimately break down the cells in the abscis- cially. NAA, however, is the only auxin regis-
sion zone. It is believed that the mode of tered for control of preharvest drop.
452 D.W. Greene
100
Control
80 AVG 90 mg l–1
Cumulative drop (%)
NAA 10 mg l–1
60
40
20
3 4 5 6 7 8 9
Time after AVG application (weeks)
Fig. 17.4. Influence of 90 mg aminoethoxyvinylglycine (AVG, ReTain®) l1 and 10 mg naphthaleneacetic

acid (NAA) l1 on preharvest drop of ‘McIntosh’/M.26 apples. AVG application was made with 0.01%
Silwet surfactant on 15 August at the calculated dilute tree-row volume (TRV). NAA application was made
as a dilute TRV application on 11 September when preharvest drop was just starting. (From D.W. Greene,
Massachusetts, unpublished data.)
17.7.2.1 Label-recommended use of NAA effects on fruit flesh firmness. The superior
performance of NAA preload over conven-
NAA is used at rates between 5 and 20 mg l1
tional application is illustrated in Fig. 17.5.
and it should be applied before significant
This approach, however, has not been effec-
drop begins. Many of the failures or poor tive for controlling drop on ‘McIntosh’ in the
response of NAA to retard drop can be attrib- north east of the USA (D.W. Greene and W.R.
uted to late application, when drop is already Autio unpublished results).
under way (Fig. 17.4). Normally 1–2 days are
required for NAA to become effective. If drop
has started, it may require up to 5 days for 17.8 Improving Fruit Appearance and
NAA to slow drop. NAA is effective for 7–12 Shape
days and a second application will be neces-
sary, therefore, to reliably retard drop after 10 Apples are just one of many choices of fruits
days. Fruit softening and reduced storage life and vegetables now available to consumers.
are likely if warm weather follows applica- The decision to purchase a product may be
tion or if harvest is delayed until ripening has spontaneous and may be based solely on
been substantially advanced. appearance and attractiveness. Therefore, any
PBR application that improves fruit appear-
ance is also likely to improve fruit sales and
17.7.2.2 Drop control using NAA preload
product movement (Looney, 1983, 1996).
Marini et al. (1993) reported that repeated
applications of NAA well in advance of the
start of preharvest drop may be more effec- 17.8.1 Improving fruit shape
tive on ‘Delicious’ than when NAA is
applied just once prior to ripening. Unrath ‘Delicious’ remains a dominant apple culti-
(1996) suggested that four weekly applica- var in the market today. A ‘Delicious’ that
tions of NAA at 5 mg l1 started 4 weeks is blocky in shape and has prominent calyx
before harvest (preloading) effectively con- lobes is perceived by the buying public to
trolled drop and it did not have adverse be preferred and have better quality. The
80
Control
NAA traditional
60
Cumulative drop (%)
NAA preload
40
20
0
1 2 3 4 5 6 7
Time after normal harvest (weeks)
Fig. 17.5. Preharvest drop of ‘Delicious’ apples following NAA application – a 10-year average between
1991 and 2000. Traditional NAA application involved 10 mg l1 at the start of harvest and a second 10 mg
l1 7 days later. Preload NAA involved application of 5 mg l1 4, 3, 2 and 1 week before the anticipated
start of harvest. (From C. Richard Unrath, North Carolina, unpublished data.)
proprietory product Promalin®, which con- price to the growers. Even if the russet is
tains equal quantities of GA4+7 and BA, is not severe, it makes fruit less attractive to
used to elongate fruit so that they have a consumers (Looney, 1993). Russeting in
‘typier’ appearance. Promalin® is applied apples is reduced by Provide®, a product
at a rate of 1.17–2.34 l ha1. If a surfactant is containing GA4 and GA7 in approximately
used, the effect of Promalin® is increased equal amounts (Greene, 1993). The label rec-
and fruit thinning is also increased. ommendation is to apply two to four sprays
Promalin® should be applied at the first of 475–950 ml ha1 at 7–10-day intervals
favourably warm weather opportunity starting at petal fall. The label restricts the
after the king flower opens. Promalin® will amount that can be applied to 99 g (a.i.)
thin young trees, so use of this product ha1. Early applications are more effective
should be restricted to use on mature than later applications, so, if only two
blocks or on young trees that are approach- sprays are applied, they should be made
ing stable production. Rates above 50 mg during the first 2 weeks after petal fall. The
l1 may result in reduced return bloom. It use of a surfactant with Provide® is not rec-
is not recommended to use NAA on ommended, since the surfactant itself may
‘Delicious’ trees that have previously been cause some russeting. Application is made
sprayed with Promalin®, since small seed- at less than TRV, frequently 935 l ha1. Since
less fruit may be produced and they fre- gibberellins inhibit flower-bud formation,
quently persist to harvest. even the relatively small amounts used to
inhibit russeting may have a negative influ-
ence on flower-bud formation for the fol-
17.8.2 Improving fruit finish lowing year.
Provide® may also suppress cracking on
Moist, humid conditions, with rain and ‘Stayman’ apples if application is made at
fluctuating temperatures, frequently result least 2–3 weeks before cracking is likely to
in fruit russeting on susceptible cultivars, begin. Application rates of 1165–2330 l ha1
such as ‘Golden Delicious’. If severe, russet per spray are made 2–3 weeks apart. The
may lower fruit grade and result in a lower addition of a surfactant is recommended.
454 D.W. Greene
17.9 Influencing Fruit Maturity and NAA is applied. Treated fruit should be
Quality monitored daily, once ripening is observed
to start. Ripening advances much more
It is to the advantage of both consumers and rapidly than on untreated trees, and over-
orchardists to have attractive, high-quality, ripening will occur if fruit are not harvested
freshly picked apples available for sale over at the appropriate time.
a longer duration than the normal harvest Ethephon-treated fruit should be treated
period for a cultivar. Several PBRs used differently following harvest. The storage life
alone or in combination can advance the and the shelf-life may be reduced. In some
harvest season by triggering early fruit years there may be no apparent effect, while
ripening or extend the normal harvest sea- in others the effect may be dramatic. Factors
son by delaying ripening. The quality of that may substantially reduce the storage life
fruit available to consumers from these of ethephon-treated fruit include using a
sources is increased. high concentration of ethephon, a long time
interval between application and harvest, the
occurrence of high temperatures during the
17.9.1 Advancing fruit ripening with time the fruit are on the tree and a long time
ethephon interval from harvest until internal fruit tem-
perature is reduced to 0°C in storage. Yield
If fruit are harvested before they are physio- will be reduced because fruit will be har-
logically mature, they will be tart, tannic, vested 1–2 weeks earlier than normal. Fruit
and starchy, lack sweetness and not have the increase in size about 1% day1 while they
characteristics typical of the cultivar. Sale of remain on the tree; thus harvesting fruit 15
inferior-quality fruit early in the season may days earlier than normal will result in a
depress future sales because consumers have potential reduction in yield of 15%.
had a bad eating experience at the beginning
of the season. Ethephon is used to advance
ripening of apples to provide a higher- 17.9.2 Increasing red colour with ethephon
quality product earlier in the season.
Ethephon should be applied between 1 Nearly all red-colouring cultivars of apples
and 3 weeks before normal harvest at con- benefit from additional colour. Furthermore,
centrations ranging from 62.5 to 125 ml per the development of suitable red colour is the
100 l water (based upon a dilute TRV appli- factor that frequently determines when a
cation). Increased red colour will be noted grower starts to harvest a block of fruit.
within 5–7 days. Flesh softening will accom- Ethephon application of 31–62 ml 100 l1
pany red colour development. The closer the water (based upon a dilute TRV application)
application is made to the time of normal made 7–10 days before anticipated harvest
harvest, the shorter the time required to may increase red colour. Improved red
observe a colour and a maturity response. colour and reduction in flesh firmness fol-
A preharvest drop strategy will be neces- lowing ethephon application is illustrated in
sary if ethephon is used. Normally 10–20 Table 17.7. This use comes with some risk,
mg NAA l1 is used for this purpose and it because there is the potential to reduce fruit
is effective for 7–12 days. The NAA can be storage life. It should be noted that ethephon
applied when the ethephon is applied or it will not completely overcome conditions
can be delayed for 3–4 days. Application of unfavourable for development of red colour.
NAA with the ethephon ensures the pres- At elevated temperatures, fruit ripen at an
ence of a drop-control compound if poor accelerated rate, while red colour may
spray weather occurs when the NAA is increase little. It is important to harvest fruit
applied. Alternatively, a delay in applica- before fruit condition is lost, and to cool the
tion is a strategy that can be used to extend fruit immediately. Good exposure of the fruit
preharvest-drop control for several addi- to light is an important component when
tional days where just one application of using ethephon. Experience has shown that
Table 17.7. Influence of preharvest application of

ethephon on 11 August on red-colour development
and flesh firmness of ‘McIntosh’ apples at harvest
on 25 August (from Greene et al., 1977).
Ethephon Red Flesh

concentration colour firmness
(mg l1) (%) (N)
0 (Control) 35c 66.0a

125 57b 63.7b
250 73a 63.3b
Values with uncommon letters are significantly

different at odds of 19 to 1.
ethephon works best on young, well-pruned A delay of ripening of 7–12 days is common
trees. Ethephon should not be used on large, in cultivars that produce large amounts of
dense trees or poorly pruned trees where ethylene, such as ‘McIntosh’. Delay in ripen-
fruit may ripen excessively before adequate ing of some other cultivars, such as ‘Gala’,
colour develops. which are characteristically low ethylene
producers, may be as long as 4 weeks.
The primary mode of action of ReTain® is
17.9.3 Delaying ripening and improving fruit to inhibit ethylene production. Conse-
quality with ReTain® quently, apples that have been treated with
ReTain® will lack many of the aromatic com-
In apple-growing areas where one cultivar ponents and flavours associated with the
predominates or where the time of ripening process of ripening. While this may not be a
of two or more cultivars overlaps, problem with some less flavourful cultivars,
orchardists are challenged to harvest the such as ‘Delicious’, this reduction in taste
fruit in a timely manner while maintaining can be very apparent and of some concern
fruit quality. In addition to retarding prehar- with new cultivars that were selected in
vest drop, application of ReTain® at 123.5 g large part because of their superior taste.
(a.i.) ha1 4 weeks before anticipated normal Some cultivars, such as ‘Delicious’,
harvest will delay ripening of fruit. All of the develop water-core (an accumulation of
application considerations mentioned earlier sorbitol in water-soaked areas within the
for the use of ReTain® use in drop control are fruit) and, if severe, this disorder can lead
also appropriate for delaying maturity. to the development of senescent break-
Ripening can be delayed by 5–14 days, down in storage. ReTain® will delay and
depending upon the weather and cultivar. reduce the amount of water-core develop-
Red-colour development, flesh-firmness loss ment. The increased firmness of fruit fol-
and the degradation of starch are all delayed. lowing ReTain® application has also been
There has been some concern expressed over demonstrated. However, the response is
the reduction in red-colour development, quite erratic and unpredictable. In some
since colour is a factor that determines fruit years firmness differences of 1 kg have
grade. However, most pomologists now agree been noted between ReTain®-treated and
that reduced red colour is a direct conse- control fruit, while in others differences are
quence of delayed ripening. When red-colour minimal. Pomologists still do not know
development is compared on fruit at a com- what causes the differences and how to
parable stage of maturity, ReTain® does not predict the response. Equally variable and
reduce red colour: it just delays development. unpredictable is the firmness response of
There are differences among cultivars fruit coming out of controlled-atmosphere
related to the time ReTain® delays ripening. storage.
456 D.W. Greene
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(eds) Tree Growth Regulators and Chemical Thinning. Washington State Cooperative Extension,
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and gibberellins A4+7. Journal of the American Society for Horticultural Science 107, 538–541.
Forshey, C.G., Elfving, D.C. and Stebbins, R.L. (1992) Training and Pruning Apple and Pear Trees. American
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cropping. Acta Horticulturae 179, 343–348.
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and Mullinex, K. (eds) Tree Fruit Physiology: Growth and Development. Good Fruit Grower, Yakima,
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bioregulants. Acta Horticulturae 329, 120–127.
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of ‘Delicious’ apples. Journal of Horticultural Science 68, 247–253.
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development in selected apple cultivars. Scientia Horticulturae 28, 355–368.
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paclobutrazol. Acta Horticulturae 179, 563–566.
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Williams, M.W. (1980) Retention of fruit firmness and increase in vegetative growth and fruit set of
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18 Diseases of Apple
Gary G. Grove1, Kenneth C. Eastwell1, Alan L. Jones2 and Turner B. Sutton3

1Irrigated Agriculture Research and Extension Center, Washington State University,
Prosser, Washington, USA; 2Department of Botany and Plant Pathology, Michigan State
University, East Lansing, Michigan, USA; 3Department of Plant Pathology, North
Carolina State University, Raleigh, North Carolina, USA

18.2 Diseases Caused by Bacteria 460
18.2.1 Fire blight 460
18.3 Diseases Caused by Fungi 468
18.3.1 Apple scab 468
18.3.2 Powdery mildew 470
18.3.3 Brown-rot diseases 472
18.3.4 Summer diseases 473
18.3.5 Phytophthora crown and root rot 476
18.3.6 European or Nectria canker 477
18.4 Postharvest Diseases 477
18.4.1 Blue mould 478
18.5 Diseases Caused by Viruses, Viroids, Phytoplasmas and Other Virus-like Agents 478
18.5.1 Chlorotic leaf spot 479
18.5.2 Apple decline (on ‘Virginia Crab’) 480
18.5.3 Flat apple 480
18.5.4 Apple mosaic 481
18.5.5 Stem pitting 481
18.5.6 Union necrosis 481
18.6 Diseases Caused by Phytoplasmas 482
18.6.1 Proliferation 482
18.6.2 Chat fruit 482
18.7 Diseases of Apple Caused by Viroids 482
18.7.1 Blister bark 482
18.7.2 Dapple apple 483
18.7.3 Dimple fruit 483
18.7.4 Fruit crinkle 483
18.8 ’Virus-like’ or Graft-transmissible Diseases of Apple with No Known Causal Agents 483
18.8.1 Green crinkle 484
18.8.2 Rubbery wood 484
18.8.3 Dead spur 485
18.9 Control Measures 485

460 G.G. Grove et al.
18.1 Introduction Blister bark has been described from South

Africa but the pathogen P. s. pv. syringae van
Apples are host to over 70 infectious dis- Hall is found as a common epiphyte on
eases, the vast majority of which are caused apple foliage worldwide. Crown gall and
by pathogenic fungi (Table 18.1). Apples are hairy root are soil-borne diseases caused by
also susceptible to diseases caused by bac- two species of Agrobacterium; crown gall is
teria, phytoplasma and virus/virus-like the most common of the two problems.
agents. The authors discuss several but not In areas where these bacterial pathogens
all economically important pre- and posthar- either are not established or are rare, bacter-
vest apple diseases in this chapter. Readers ial diseases are often avoided by planting
should keep in mind that some of the dis- pathogen-free nursery stock. Where these
eases discussed might be major in some diseases are established, they are managed
areas and minor in others. Others (e.g. fire through orchard sanitation, wound mini-
blight) are of importance or potential impor- mization, the use of resistant rootstocks and
tance wherever apples are grown. In most cultivars, site selection, cultural practices
apple-producing regions, disease control is a that promote air movement and drying of
major annual expense for the grower. For plant surfaces and, in some cases, the use of
example, in the eastern USA, the manage- copper and antibiotic sprays.
ment of apple scab can require eight to ten
protective fungicide applications annually. In
those areas, the apple grower must manage 18.2.1 Fire blight
early-season diseases, such as apple scab and
cedar apple rust, as well as a group of dis- Fire blight is of major concern in all countries
eases termed ‘summer diseases’. Conversely, where apples are grown including countries
in the arid production regions of the Pacific that are currently free of this disease
Northwest, these diseases are non-existent or (Vanneste, 2000). When fire blight is epi-
sporadic in nature and powdery mildew is demic, it can cause serious tree loss in nurs-
much more problematic. eries and orchards (Plate 18.1), even leading
Successful disease management usually to orchard removal. In high-risk areas, fire
results from the integration of several meth- blight is limiting the planting of some highly
ods of disease control. The use of resistant susceptible apple cultivars and rootstocks.
rootstocks and scions, fungicides, bacteri- Strict quarantines and restrictions are main-
cides, biological control agents, environmen- tained in countries where the disease does
tal modification and site selection are some not currently occur; enforcing quarantine
of the means used to control apple diseases. regulations and restrictions and, if necessary,
The precise combination and order of control eradicating the disease are very costly.
measures are usually disease-specific. The fire blight pathogen kills fruit-bear-
ing spurs, branches and entire trees. Infected
blossoms are initially water-soaked and
18.2 Diseases Caused by Bacteria darker green; spurs with infected blossoms
(Table 18.2) turn brown to dark brown and collapse after
4–5 days. Infected shoots turn brown to
Fire blight, blister spot, blister bark, crown black from the tip and bend near the tip to
gall and hairy root are diseases of apple resemble a shepherd’s crook (Plate 18.2).
caused by bacteria. Fire blight is the most When shoots are invaded from the base, the
important of these diseases and is described basal leaves and stem turn brown to black.
in more detail below. Blister spot, caused by Leaves may exhibit discoloration of the
Pseudomonas syringae pv. papulans (Rose) midrib, followed shortly by a darkening of
Dhanvantari, occurs in North America and the lateral veins and surrounding tissues.
Europe. It causes a fruit spot on Bark on infected branches and scaffold
‘Mutsu’/‘Crispin’, ‘Fuji’, ‘Redcort’, ‘Sun limbs is darker than normal. When the outer
Crisp’, ‘Smoothee’ and a few other cultivars. bark is peeled away, the inner tissues are
Apples - Chap 18
Table 18.1. Fungal diseases of apples, names of the asexual and sexual stages of the causal fungi and their reproductive stages, the symptoms and damage
most likely to be encountered and general methods of control. (Malus domestica Borkh.) A.L. Jones, primary collator (last update 3/12/93).
Disease (distributiona) Fungal pathogen Reproductive stage Common symptomsb Main control method
21/3/03
Alternaria blotch (A, Alternaria mali Roberts; = Alternaria Conidia/conidiophores Leaf spots, defoliation Fungicides,
AF, NA) alternata (Fr.:Fr) Kessl apple pathotype cultivar selection
Alternaria rot (W) Alternaria alternata (Fr.:Fr.) Keissl Conidia/conidiophores Decay of fruit None
3:03 pm
American brown rot Monilinia fructicola (G. Wint.) Honey Conidia in sporodochia; Decay of fruit (P) Avoid preharvest injury to fruit
(A, AF, NA,O) ascospores in apothecia
Anthracnose canker Pezicula malicorticis (H. Jacks.) Nannf Conidia in acervuli; Cankers on wood, decay Fungicides
and bull’s-eye rot (E, Cryptosporiopsis curvispora (Peck) ascospores in apothecia of fruit (P)
NA, O) Gremmen in Boerema & Gremmen
Page 461
[anamorph]
Apple scab (W) Venturia inaequalis (Cooke) G. Wint. Ascospores in pseudothecia; Leaf spots, scabs on fruit, Sanitation, fungicides, resistant
Diseases of Apple
Spilocaea pomi Fr.:Fr. [anamorph] conidia/conidiophores defoliation cultivars
Apple ring rot and Botryosphaeria berengeriana De Not. (syn. Conidia in pycnidia; Cankers on wood, decay Fungicides
canker (A) Physalospora piricola Nose) ascospores in pseudothecia of fruit
Armillaria (shoestring) Armillaria mellea (Vahl:Fr.) P. Kumm. Basidiospores in basidiocarps Decay of roots None, avoid problem sites
root rot (W) (mushrooms); rhizomorphs
Bitter rot (W) Glomerella cingulata (Stoneman) Spauld. Conidia in acervuli; Decay of fruit Fungicides, sanitation
& H. Schrenk ascospores in perithecia
Colletotrichum gloeosporioides (Penz.)
Penz. & Sacc. in Penz. [anamorph]
Colletotrichum acutatum J.H. Simmons
Black pox (NA) Helminthosporium papulosum Berg. Conidia/conidiophores Spots on fruit and leaves Fungicides
Black root rot (NA for Xylaria mali Fromme Ascospores in perithecia Decay of roots None
X. mali; W) Xylaria polymorpha (Pers.:Fr.) Grev.
Black rot, frog-eye Botryosphaeria obtusa (Schwein.) Shoemaker Conidia in pycnidia, Spots on leaves and fruit, Sanitation, fungicides
leaf spot and canker Sphaeropsis malorum Berk. [anamorph] ascospores in pseudothecia defoliation, cankers on
(AF, E, NA, SA, O) wood
Blister canker (nail- Biscogniauxia marginata (Fr.) Pouzar Conidia in a sclerotia-like Cankers on branches Removal of infected branches
head canker) (NA) = Nummularia discreta (Schwein.) Tul. & C. stroma, ascospores in
Tul. perithecia
461
Continued
Apples - Chap 18
462
Blotch (NA) Phyllosticta solitaria Ellis & Everh. Conidia in pycnidia Blotches on fruit, leaf Fungicides
21/3/03
spots, blisters in bark
Blue mould (W) Penicillium spp. Conidia/conidiophores Decay of fruit (P) Packing-house sanitation,
P. expansum Link fungicides
Brooks fruit spot (NA) Mycosphaerella pomi (Pass.) Lindau Conidia/conidiophores; Spots on fruit, often at Fungicides
3:03 pm
Cylindrosporium pomi C. Brooks ascospores in pseudothecia calyx end
[anamorph]
Brown-rot blossom Monilinia laxa (Aderh. & Ruhl.) Honey Conidia in sporodochia Blighting of blossoms, Fungicides
blight (A, AF, E, NA, spur dieback
Page 462
SA)
Calyx-end rot (NA) Sclerotinia sclerotiorum (Lib.) de Bary Ascospores in apothecium Decay of fruit None
G.G. Grove et al.

Clitocybe root rot (NA) Armillaria tabescens (Scop.) Dennis et al Basidiospores in basidiocarps Decay of roots None
= Clitocybe tabescens (Scop.) Bres.
Diaporthe canker (A) Diaporthe tanakae Kobayashi & Sakuma Conidia ( and ß spores) in Cankers on 1- and Sanitation
Phomopsis tanakae Kobayashi & Sakuma pycnidia; ascospores in 2-year-old shoots
[anamorph] perithecia
Diplodia canker (E, Botryosphaeria stevensii Shoemaker Pycnidia and pseudothecia; Cankers on branches Pruning out of infected
NA, O) = Physalospora malorum Shear et al. often in the same stroma branches
Diplodia mutila (Fr.: Fr.) Mont. [anamorph]
European brown rot Monilinia fructigena Honey in Whetzel Conidia in sporodochia; Blossoms and spur Sanitation, fungicides
(A, E, AF) Monilia fructigena Pers.:Fr. [anamorph] ascospores in apothecia blight, decay of injured
Monilinia laxa (Aderhold & Ruhland) fruit
Honey
Fish-eye rot (A, E, Butlerelfia eustacei Weresub & Illman Basidiomycete, may produce Decay of fruit (P) None
NA) = Corticium centrifugum (Lév.) Bres. basidiospores in culture
Fly-speck (W) Schizothyrium pomi (Mont.:Fr.) Arx Conidia in pycnidia Small, superficial, dark Cultural practices, fungicides
Zygophiala jamaicensis E. Mason spots on fruit
[anamorph]
Glomerella leaf spot Glomerella cingulata (Stoneman) Spauld. & Conidia in acervuli; Spots on leaves, Fungicides
(NA, SA) H. Schrenk ascospores in perithecia defoliation
Colletotrichum gloeosporioides (Penz.)
Penz. & Sacc. in Penz. [anamorph]
Apples - Chap 18
Grey-mould rot (W) Botrytis cinerea Pers. Fr. Conidia and sclerotia (rare in Decay often at calyx end Packing-house sanitation and
= dry-eye rot, Botryotinia fuckeliana (de Bary) Whetzel nature) of fruit; nest rot of fruit in fungicides
blossom-end rot [teleomorph] storage (P)
Leptosphaeria canker Diapleella coniothyrium (Fuckel) Barr Leaves
21/3/03
and fruit rot (W) = Leptosphaeria coniothyrium (Fuckel) Sacc.
Coniothyrium fuckelii Sacc. [anamorph]
Leucostoma canker Leucostoma cincta (Fr.:Fr.) Hohn. Perithecial stroma with a Cankers on branches Removal of affected branches
and dieback (E, NA) Cytospora cincta Sacc. [anamorph] central pycnidium
3:03 pm
Valsa auerswaldii Nitschke
= Leucostoma auerswaldii (Nitschke) Hohn.
Cytospora personata Fr. [anamorph]
Marssonina blotch (A, Diplocarpon mali Harada & Sawamura Ascospores in apothecia; Leaf spots, defoliation, Sanitation, fungicides
Page 463
NA, E) Marssonina coronaria (Ellis & J.J. Davis) conidia in acervuli spots on surface of fruit
J.J. Davis [anamorph]
Mouldy core and core Alternaria spp.; Stemphylium spp.; Conidia of various types Discoloration and decay Packing-house sanitation
Diseases of Apple
rot (W) Cladosporium spp.; Ulocladium spp. around the fruit core
Epicoccum spp.; Coniothyrium spp.
and Pleospora herbarum (Pers.) Rabenh.
wet core rot, mainly Penicillium spp.
Monilia leaf blight (A) Monilinia mali (Takahashi) Whetzel Ascospores in apothecia; Blossom and spur blight, Sanitation and chemical control
Monilia spp. [anamorph] conidia with disjunctors decay of young fruit
Monochaetia twig Seiridium unicorne (Cooke & Ellis) Sutton Cankers in wounds on None
canker (NA) = Monochaetia mali (Ellis & Everh.) Sacc. small twigs
Lepteutypa cupressi (Nattras et al.) H.J.
Swart [teleomorph]
Mucor rot (AF, NA, Mucor spp. Sporangiospores and Soft, watery decay of Packing-house sanitation
O) M. piriformis E. Fischer zygospores fruit (P)
Nectria canker (A, AF, Nectria galligena Bres. in Strass. Conidia on phialides; Cankers on branches, Fungicides
NA, SA, O) Cylindrocarpon heteronemum (Berk. & ascospores in perithecia occasional fruit decay
Broome) Wollenweb. [anamorph]
Nectria twig blight Nectria cinnabarina (Tode:Fr.) Fr. Conidia on sporodochia; Cankers on branches Removal of affected branches
= coral spot (E, O, NA) Tubercularia vulgaris Tode:Fr. [anamorph] ascospores in perithecia and trunk
Continued
463
Apples - Chap 18
464
Peniophora root canker Peniophora sacrata G.H. Cunn. Basidiomycete with crust-like Infected roots with Resistant rootstocks and
21/3/03
(O) fruiting-bodies characteristic cracks sanitation
Perennial canker (NA) Neofabrae perennans Kienholz Conidia in acervuli; Cankers on branches, Sanitation, fungicides
Cryptosporiopsis perennans (Zeller & ascospores in apothecia decay of fruit
Childs) Wollenweb. [anamorph]
3:03 pm
Phomopsis canker, fruit Phomopsis mali Roberts Conidia ( and ß spores) in Cankers on wood, fruit None
decay and rough bark Diaporthe perniciosa Em. Marchal pycnidia, ascospores in decay (P)
(A, E, NA) [teleomorph] perithecia
Phymatotrichum root Phymatotrichopsis omnivora (Duggar) Hyphae with conidia on Decay of roots None
Page 464
rot Hennebert conidiophores and sclerotia
= cotton root rot (NA) = Phymatotrichum omnivorum Duggar
Phytophthora crown, Phytophthora spp. Zoospores produced in Decay of crown and root Cultural practices, host
G.G. Grove et al.

collar and root rot; P. cactorum (Lebert & Cohn) J. Schröt. sporangia from hyphae or tissues resistance, chemical treatment
(sprinkler rot) (W) P. cambivora (Petr.) Buisman germinating oospores or
P. cryptogea Pethybr. & Lafferty chlamydospores
P. megasperma Drechs.
P. syringae (Kleb.) Kleb.
Phytophthora fruit rot Phytophthora cactorum (Lebert & Cohn) See above Rotting of fruit Late-season fungicide sprays
(E, NA) J. Schröt.; or postharvest treatments
Phytophthora syringae (Kleb.) Kleb.
Pink mould rot (W) Trichothecium roseum (Pers.:Fr.) Link Conidia Decay of fruit (P) Prevented by refrigerated cold
= Cephalothecium roseum Corda storage
Powdery mildew (W) Podosphaera leucotricha (Ellis & Everh.) Conidia/conidiophore; Powdery spots on leaves, Cultivar selection, fungicides
E.S. Salmon ascospores in cleistothecia fruit russeting
Rosellinia root rot (W) Rosellinia necatrix Prill. Ascospores in perithecia; Decay of roots Cultural practices, solarization
= Dematophora root rot Dematophora necatrix R. Hartig pycnidia on synnemata
[anamorph]
Rust, American Gymnosporangium globosum (Farl.) Farl. Pycnia and aecidia on apple; Leaves Removal of alternative host,
hawthorne (NA) telentospores in telial horns fungicides
on cedar
Rust, cedar apple (NA) Gymnosporangium juniperi-virginianae Pycnia and aecidia on apple; Spots on leaves, Removal of alternative host,
Schwein telentospores in telial horns defoliation and distortion fungicides
on cedar of fruit
Apples - Chap 18
Rust, Japanese apple Gymnosporangium yamadae Miyabe ex Pycnia and aecidia on apple; Leaves Removal of alternate host,
(A) Yamada telentospores in telia on fungicides
Juniperus chinensis
Rust, Pacific Coast Gymnosporangium libocedri (C.Henn.) F. Aecidia on apple;
21/3/03
pear (NA) Kern telentospores in telia on
Libocedrus decurrens
Rust, quince (NA) Gymnosporangium clavipes (Cooke & Peck) Pycnia and aecidia on apple; Distortion of fruit Removal of alternative host,
Cooke & Peck in Peck telentospores in telia on cedar fungicides
3:03 pm
Side rot (E, NA) Phialophora malorum (M.N. Kidd & A. Conidia at the apex of Decay of fruit (P) Cultural practices, postharvest
Beaumont) McColloch phialides treatments
Silver leaf (W) Chondrostereum purpureum (Pers.:Fr.) Pouzar Basidiospores Wood decay Cultural practices
Sooty-blotch complex Peltaster fructicola (Johnson, Sutton, Conidia in pycnidia Blotches of various size Cultural practices and
Page 465
(W) Hodges); Geastrumia polystigmatis on the surface of fruit fungicides
Batista & M.L. Farr; Leptodontium
elatius (G. Mangenot) (De Hoog); and
Gloeodes pomigena (Schwein.) Colby)
Diseases of Apple
Southern blight (AF, Sclerotium rolfsii Sacc. Sclerotia Decay at base of trunk Cultural practices
NA, SA) Athelia rolfsii (Curzi) Tu & Kimbrough
[teleomorph]
Thread blight Corticium stevensii Burt Rhizomorphs and sclerotia; Blighting of branches Fungicides
= Hypochnus leaf blight = Pellicularia koleroga Cooke basidia with basidiospores
(NA) = Hypochnus ochroleucus Noack
Valsa canker(A) Valsa ceratosperma (Tode:Fr.) Maire Pycnidia and perithecia in a Cankers on branches Sanitation
Cytospora sacculus (Schwein.) Gvritischvili stroma
[anamorph]
Violet root rot (O) Helicobasidium mompa Tanaka Basidiospores in basidiocarps Root decay Soil sterilization
White root rot (NA) Scytinostroma galactinum (Fr.) Donk Basidia on a resupinate Decay of roots None
= Corticium galactinum (Fr.) Burt hymenium
White rot (AF, NA, Botryosphaeria dothidea (Moug.) Ces. & De Conidia in pycnidia; Decay of fruit, cankers Fungicides and sanitation
SA, O) Not. ascospores in pseudothecia on limbs and twigs
Fusicoccum aesculi Corda [anamorph]
Zonate leaf spot (A) Cristulariella moricola (Hino) Redhead Ascospores in apothecia; Leaf spot, defoliation Sanitation
Grovesinia pyramidalis M. Cline et al. conidia
[teleomorph]
aApproximate
465
geographical distribution of the disease: A, Asia; AF, Africa; E, Europe; NA, North America; O, Oceania; SA, South America; W, worldwide.
bP, postharvest disease problem.
Table 18.2. Bacterial diseases of apple, scientific name of the pathogen, symptoms and main control method.
Disease Common
(distributiona) Pathogen symptoms Main control method
Fire blight (E, NA, Erwinia amylovora Blight, cankers, Resistant cultivars and
New Zealand) dead trees rootstocks, sanitation,
antibiotics
Blister spot (E, NA) Pseudomonas syringae pv. Spots on fruit Avoiding susceptible
papulans (Rose) Dhanvantari cultivars, antibiotics
Blister bark (AF) Pseudomonas syringae pv. Terminal dieback, None
syringae van Hall blossom blight,
Crown gall (W) Agrobacterium tumefaciens Galls on roots Sanitation, biocontrol but
(E.F. Smith & Townsend) Conn often less effective than
on Prunus species
Hairy root (W) Agrobacterium rhizogenes Excessive production Sanitation
(Riker et al.) Conn of fibrous roots
aApproximate geographical distribution of the pathogen: A, Africa; E, Europe; NA, North America, W,
worldwide.
water-soaked with reddish streaks when and LB media (see Bereswill et al., 1998, for
first invaded; later the tissues are brown. As the composition of these media), patho-
lesion expansion slows, the bark sometimes genicity tests performed on immature fruit,
cracks, delineating a canker. Infected fruit seedlings or vigorous plants, and molecular
develop a brown-to-black decay. During assays (Bereswill et al., 1995; McManus and
wet, humid weather, droplets of whitish to Jones, 1995).
reddish-brown, sticky liquid (known as The pathogen overwinters in cankers
ooze) seeps from the surface of infected tis- located on branches and tree trunks (Fig.
sues (Plate 18.3). Infected rootstocks may 18.1). Ooze can begin to appear on the sur-
show bleeding or ooze from rootstock tissue face of these cankers at or just before the
early in summer and internal necrosis of the onset of bloom. Bacteria are disseminated to
bark and the leaves of the scion turn reddish flowers by splashing rain and, occasionally,
early in the autumn. flies and other insects that visit both bacterial
Fire blight is caused by Erwinia amylovora ooze and blossoms. Eventually, honey bees
(Burrill) Winslow et al., a Gram-negative, visit infected blossoms and pick up pollen or
rod-shaped, non-fluorescent bacterium with nectar contaminated with bacteria. Spread of
peritrichous flagella. More than 130 species bacteria from flower to flower by bees is
in 39 genera of the Rosaceae are hosts. rapid. Bacteria colonize the stigmatic surface
Important hosts include apple, pear, orna- of the pistils of healthy apple and pear flow-
mental Pyrus and Malus species, quince ers, resulting in epiphytic populations of
(Cydonia oblonga), loquat (Eriobotrya japon- bacteria (Thomson, 1986). Climatic condi-
ica), hawthorn (Crataegus spp.), Cotoneaster tions govern the rate of spread and severity
spp., Sorbus spp., Pyracantha spp. and Rubus of blossom blight and even the infection
spp. Most strains of E. amylovora from Rubus process itself. Temperatures between 18.3
do not infect apple and are therefore consid- and 30°C (accompanied by rain or high
ered to be pathologically distinct. Primary humidity during the day) favour infection.
isolation of the bacteria is by culturing on Although bacteria invade the flowers pri-
Luria–Bertani (LB) agar or some semi-selec- marily through natural openings, storms
tive medium. They are distinguished from containing wind-driven rain or hail are
other plant pathogenic bacteria based on important in spreading bacteria during the
colony colour and morphology on MM2Cu summer months, which under some condi-
Diseases of Apple 467
Fig. 18.1. The fire blight disease cycle. The pathogen overwinters in cankers. At the onset of bloom, bacteria
begin to ooze on to the surface of cankers; splashing rain, along with flies and other insects, moves the
bacteria to open flowers. Then, honey bees spread the bacteria from flower to flower on contaminated
pollen and nectar. The bacteria multiply on the stigmatic surface of the pistils to produce high epiphytic
populations. Favourable combinations of moisture and temperature govern infection of the flowers. Summer
storms with wind-driven rain spread the bacteria, leading to sudden and severe outbreaks of diseases.
Inoculum for secondary infection originates from droplets of ooze produced on infected flowers, fruit and
shoots. (Figure courtesy A.L. Jones.)
tions lead to sudden and severe outbreaks of to prevent the importation of plant material
disease. Inoculum for secondary infection that may harbour the pathogen.
originates from droplets of ooze produced In countries where fire blight is a prob-
on infected flowers, fruits and shoots. lem, sanitation – the removal of infected por-
Temperature, as measured by the accumu- tions of the tree – is critical to the success of
lation of degree-days (DD), governs the rate other control measures and is effective pro-
of symptom development. Symptoms are vided it is done during the early stages of an
expressed when 55 DD (base 12.5°C) are epidemic. New orchards should be planted
accumulated following the infection date. on blight-resistant rootstocks if available. A
The susceptible rootstocks M.9 and M.26 are realistic plan for controlling fire blight is
infected by systemic movement of bacteria needed before making large plantings of
through symptomless scion tissue or through blight-susceptible cultivars. Antibiotics have
infected rootstock shoots (Momol et al., 1998). been highly effective for preventing the blos-
Although the rootstock Bud.9 is blight-sus- som-blight stage of fire blight, but in some
ceptible, rootstock blight has not developed regions or orchards control with antibiotics is
on many Bud.9/scion combinations. Losses no longer possible, due to the development
from rootstock blight can be avoided by of streptomycin-resistant strains of E.
using resistant rootstocks, such as G.16, G.30, amylovora. Predictive models, particularly
G.65 and other blight-resistant rootstocks Maryblyt and Cougarblight in the USA and
from the US Department of Agriculture– Billings’ integrated system and Firescreens in
Agricultural Research Service (USDA- Europe, help growers identify potential
ARS)/Cornell apple-rootstock programme. infection periods (Billings, 2000). Such infor-
Many practices help to prevent or reduce mation is helpful in timing antibiotic treat-
the severity of fire blight. In countries where ment and for avoiding unnecessary
E. amylovora is not established, only trees pro- treatment. Bacterial antagonists that sup-
duced in fire-blight-free regions should be press fire blight may be integrated with
used when establishing new orchards. These antibiotics to control flower infections
countries may already have strict quarantines (Stockwell et al., 1996).
18.3 Diseases Caused by Fungi symptoms are found on both sides of leaves.
Conidia are produced abundantly in new
Within this diverse group of plant pathogens lesions; therefore, lesions appear as velvety
are the causes of most apple diseases (Table brown to olive spots that turn black with age
18.1). Fungi pathogenic to apple can cause (Plate 18.4). Severe infection can cause leaves
root rots, leaf spots, leaf blights, blossom to abscise, resulting in defoliated trees. Return
blights, fruit decay, fruit spots, defoliation bloom on trees defoliated in midsummer is
and trunk, branch and twig cankers. The often reduced due to a lack of flower-bud for-
fungal diseases discussed in this chapter are mation the previous summer. Fruit infections
caused by fungi belonging to the general tax- resemble leaf infections when young but turn
onomic groups represented by the mush- brown and corky with age (Plate 18.5). Early-
rooms (Basidiomycetes) and morels season infections are often associated with the
(Ascomycetes). The phylum Oomycota also blossom end of the fruit; later they can occur
contains the water moulds, an important anywhere on the surface. Scab infections
group of plant pathogens formerly classified result in uneven growth of fruit and even
as fungi. Fungi in the ascomycete group cracking of the skin and flesh. Rough, black
have scientific names for the anamorph and circular lesions develop in cold storage on
teleomorph. The anamorph is the asexual fruit infected close to harvest. The latter phase
form (also called the imperfect stage) of the is known as pinpoint scab. Infection of blos-
fungus and produces asexual spores (such as soms and bud scales may be observed in
conidia) or no asexual spores. The teleo- high-inoculum orchards when infection peri-
morph is the sexual form (also called the per- ods occur at critical times. Lesions on blos-
fect or sexual state) and produces sexual soms and bud scales resemble those on leaves
spores (ascospores) in various fruiting bodies but are seldom observed because these tissues
(pseudothecia, perithecia, apothecia, etc.). normally drop to the ground before the symp-
These fungi generally reproduce by spores toms are well developed.
that are dispersed by air currents or splash- Venturia inaequalis (Cooke) G. Wint.,
ing water. Fungal diseases are prevented or anamorph Spilocea pome Fr., causes apple
managed by using resistant cultivars and scab. It has one sexual cycle and a series of
rootstocks, site selection, sanitation, cultural asexual cycles per year. Flask-shaped, ascus-
practices and fungicide practices. Specific bearing fruiting bodies (pseudothecia)
management measures vary according to develop in overwintering infected leaves.
disease and geographical location. Pseudothecia form as a result of the interac-
tion of two strains of opposite mating types
(heterothallic fungus). Asci in pseudothecia
18.3.1 Apple scab are eight-spored. Mature ascospores are two-
celled, with the upper cell shorter and wider
Scab occurs in virtually all apple-producing than the lower cell, yellowish green to tan
regions worldwide (MacHardy, 1996). The and with smooth walls. Conidia are one-
disease is an annual threat in cool, humid celled, yellowish-olive and pointed.
regions with frequent rainfall in spring and From late autumn to spring, microscopic,
early summer. In semiarid regions, scab is a black, pimple-like pseudothecia develop in
threat in years with above-normal rainfall or leaves on the orchard floor (Fig. 18.2).
in orchards where artificial wetting periods Normally, pseudothecia contain mature
are created from the improper use of over- ascospores when the blossom buds start to
head irrigation. open in spring. Maturation and discharge of
Scab attacks the leaves and fruit through- ascospores lasts about 5–9 weeks. When
out most of the growing season; blossoms and leaves on the orchard floor become wet from
bud scales are attacked for short periods in rain, spores are ejected into the air. Air cur-
spring and late summer, respectively. rents carry them to the emerging tissues,
Symptoms first appear on the undersides of where infection occurs. Young leaves and fruit
leaves, the side exposed as buds open. Later, are highly susceptible to infection, but their
Fig. 18.2. The apple scab disease cycle. The fungus Venturia inaequalis overwinters in leaves on the orchard
floor and sometimes in apple buds. Pseudothecia with ascospore-bearing asci in dead leaves discharge their
ascospores when the leaves become wet from rain. Ascospores produce the first, or primary, infections on
the young fruit and leaves. Conidia produced on the inner surface of apple-bud scales can also cause
primary infections. Depending on temperature, 9–17 days are required before new scab lesions appear on
the infected tissues. Conidia, formed in continual crops of new lesions, produce secondary infections.
Secondary cycles are repeated many times until leaf fall in autumn. (Figure courtesy A.L. Jones.)
susceptibility declines with maturity. Spore but fails to develop until after the fruit has
germination begins soon after ascospores been held in cold storage for several months.
land on wet leaves or fruit. Prevailing temper- Infections can also build up in leaves after
atures govern the infection rate, provided a harvest and prior to normal leaf abscission.
continuous film of moisture is present for ger- The fungus overwinters in these leaves and
mination of the spores. Depending on the they are often the source of very high levels
average temperature after penetration, 9–17 of inoculum the following spring.
days are required before the appearance of Basic studies on the biology and epidemi-
the olive-green, velvety scab lesions. In some ology of apple scab from the 1920s to the
apple-producing regions and on some culti- 1940s established a rational basis for scab
vars, the fungus also overwinters in lesions control and these concepts continue to be
on twigs and bud scales (Becker et al., 1992); refined. The initial ascosporic inoculum is
conidia produced in these lesions are a second usually present in large amounts; therefore,
form of primary inoculum. However, the estimating the risk of the initial ascosporic
number of conidia available from overwinter- inoculum and forecasts of its efficiency are
ing lesions at bud break is low compared with very important for determining when to ini-
the number of ascospores potentially avail- tiate scab-control programmes and the
able from leaf litter. application frequency. Inoculum risk is
Secondary infections are initiated by coni- determined by monitoring pseudothecia
dia produced in primary and secondary from early spring to midsummer to deter-
lesions. Conidia can be produced in new mine whether ascospores are mature and
lesions beginning as soon as 7–9 days after available for discharge and by detecting
infection and these lesions can produce coni- ascospore release in orchards during wetting
dia in continuous crops for several weeks. periods. Statistical models have been devel-
Conidial germination and infection occur oped to predict ascospore maturity based
under about the same conditions as germina- on DD accumulations and these models
tion and infection by ascospores. Secondary are replacing ascospore-monitoring pro-
infection on fruit can occur in the autumn grammes. Inoculum risk forecasts assume
high inoculum levels, an assumption that is fungal antagonists to the foliage just prior to
usually incorrect for orchards where scab leaf fall (Carisse et al., 2000), but this strategy
was well controlled the previous season. alone is not adequate for season-long scab
Therefore, methods for assessing differences control. This approach may be more feasible
in inoculum among orchards based on a in areas with lower amounts of overwinter-
potential ascospore density (PAD) system ing inoculum and mild winters.
were developed (Gadoury and MacHardy, Scab is controlled primarily with fungi-
1986). It was found that in low-inoculum cides applied in predetermined schedules,
orchards spray programmes in spring could beginning at green tip. The fungicides are
be delayed until about the tight-cluster stage applied on a 7–14-day interval with eight to
of bud development, rather than green tip in ten applications per season. Several classes
the normal spray schedule for high-inocu- of fungicides are available for apple-scab
lum orchards. Integrating inoculum risk control. They are often rotated during the
with environmental risk is accomplished by season or applied as mixtures because of the
the identification of ‘infection periods’. high potential of the scab fungus to develop
Rainfall is necessary for the discharge of fungicide-resistant strains.
ascospores – free water for the germination
of ascospores and penetration of tissues. The
rate of this infection process is temperature- 18.3.2 Powdery mildew
dependent, and the duration of wetness
required for successful infection across a Powdery mildew is circumglobal and a
wide range of temperatures is well known potentially serious disease wherever apples
(Jones, 1998). Therefore, temperature and are grown. The disease is particularly prob-
wetness measurement can be used to fore- lematic in the western and mid-Atlantic
cast the success or failure of each ascospore- regions of the USA. Severe infections reduce
discharge event. These events can be tree photosynthesis and transpiration (Ellis
monitored with environmental sensors et al., 1981) and may result in partial defoli-
linked to small computers placed in the ation (Ogawa and English, 1991). Chronic
orchard or with automated weather stations infections can result in reduced tree vigour,
connected directly to computers. Fungicides poor return bloom and yield reduction.
with post-infection efficacy are applied after Fruit infection can result in the rejection of
predicted but unprotected infection periods. fruit for fresh-market sale. Apple cultivars
Apple-breeding programmes aiming for vary in their susceptibility to powdery
high-quality, disease-resistant cultivars are in mildew. ‘Gala’, ‘Braeburn’, ‘Ginger Gold’,
progress in New Zealand and some ‘Mutsu’ (Crispin), ‘Cortland’, ‘Baldwin’,
European and North American countries. ‘Monroe’, ‘Idared’, ‘Honeycrisp’, ‘Jonathan’,
Over 50 scab-resistant cultivars have been ‘Granny Smith’, ‘Ginger Gold’, ‘Rome
released and are gaining in commercial Beauty’, ‘Jonagold’ and ‘Pink Lady®’ are
acceptance as fruit quality and other horti- particularly susceptible. ‘Delicious’ and
cultural characteristics are improved. ‘Golden Delicious’ are less susceptible.
Guarding against races of the scab pathogen Leaves, flowers and fruit are susceptible
that can overcome the resistance sources to infection by the powdery mildew fungus.
used by apple breeders is a high priority in The foliage of new terminal growth is
these breeding programmes (Bénaouf and extremely susceptible to infection. The ini-
Parisi, 2000). tial signs of powdery mildew consist of
Prevention of pseudothecia formation in white to grey felt-like patches on the lower
overwintering leaves would probably elimi- leaf surface. These patches are comprised of
nate scab. Unfortunately, complete elimina- masses of fungal mycelia and spores (coni-
tion of pseudothecia is not possible under dia). As disease progresses, mildew signs
orchard conditions using current methods. may also appear on the upper leaf surface
Spring ascospore production can be reduced and eventually cover the entire leaf. Infected
by making autumn applications of urea or foliage may curl, blister and eventually
become brittle and necrotic (Plate 18.6). are therefore primary inocula, which initiate
Internodal shortening may occur on additional new infections. This cycle of
severely infected shoots. Infected flower sporulation and foliar infection can continue
petals are distorted, stunted and light yel- as long as susceptible foliage is being pro-
low to light green. Fruit infection typically duced. This secondary phase of powdery
results in stunting accompanied by the pres- mildew can cycle many times during the
ence of a fine network of rough lines (russet- growing season. Temperature is the most
ing) (Plate 18.7). important factor affecting disease develop-
Powdery mildew is caused by Podosphaera ment. Conidia germinate at temperature
leucotricha (Ell. & Ev.) E.S. Salmon. The fun- between 10 and 25ºC; the optimum tempera-
gus is superficial on the host surface but tures for germination are 20–22ºC. The sex-
withdraws water and nutrients from host tis- ual stage (ascocarps) of P. leucotricha
sue, using a structure called a haustorium. occasionally forms on infected twigs.
The fungus produces barrel-shaped conidia Ascocarps are minute, brown to black, spher-
in chains, which are dispersed by air cur- ical fruiting bodies that contain eight unicel-
rents and initiate subsequent infections of lular ascospores. Various researchers have
foliage and fruit (Fig. 18.3). P. leucotricha suggested that this stage is insignificant in
survives through winter as mycelium in the epidemiology of the disease. Fruits are
infected buds. This mode of perennation especially susceptible to infection during the
results in the production of ‘flag shoots’ bloom period (Daines et al., 1984).
(shoots covered with powdery mildew) Powdery-mildew epidemics are favoured
when shoots emerge in the spring. Winter by high humidity; therefore problems with
temperatures are the most important factor mildew can often be avoided or reduced
affecting the amount of carry-over inoculum. with cultural practices that promote air
Temperatures colder than 12ºC kill the fun- movement and light penetration. On suscep-
gal mycelium in buds; temperatures lower tible cultivars, effective disease management
than 24ºC may kill the infected buds usually depends on a fungicide-spray pro-
(Spotts et al., 1981). Conidia produced on flag gramme. Benzimidazole, sulphur, horticul-
shoots initiate the first spring infections and tural mineral, oils, demethylation-inhibiting
Fig. 18.3. The apple powdery mildew disease cycle. The fungus overwinters as mycelium in infected buds.
Mildew conidia, produced on leaves arising from the infected buds, become wind-borne and cause primary
infections on healthy leaves and fruit. Secondary infections are produced from conidia produced in both
primary and other secondary infections and are repeated until shoot growth stops in late summer. (Figure
courtesy A.L. Jones.)
(DMI) and strobilurin fungicides are effec- pathogen present. M. laxa causes blossom
tive against powdery mildew. Spray pro- blight, spur dieback and cankering of
grammes should commence at tight cluster branches; M. fructigena causes fruit rot and
and continue until the production of new sometimes cankers when the fungus
foliage ceases. Because powdery mildews spreads into branches from the fruit; and
can develop resistance to benzimidazole, M. fructicola causes a fruit rot. Monilia leaf
DMI and strobilurin fungicides, rational blight infects young apple leaves;
resistance-management strategies usually mycelium invading from leaves kills flower
require the inclusion of two or more fungi- clusters, young fruits and fruiting spurs.
cide classes in a season-long programme. The species of Monilinia can be differenti-
DMI fungicides applied in apple-scab-man- ated by microscopic observation of conidia
agement programmes generally provide the formed in chains in culture or on infected tis-
added benefit of mildew control. sue. M. mali can be differentiated by disjunc-
A predictive system has been developed tors present within the conidial chains. M.
for aid in managing infections caused by P. fructicola, M. fructigena and M. laxa can be
leucotricha. Podem© (Xu, 1999) is a system differentiated based on cultural characteris-
developed in the UK that simulates epi- tics, isozyme variation, vegetative interac-
demics of secondary mildew on vegetative tions and PCR assays.
shoots. The effects of weather on conidial The life cycles of these pathogens differ
production, dispersal and germination are only in the role that the perfect stage
used to calculate a favourability index. The (apothecia) plays in the overwintering of the
model itself is driven by hourly ambient fungi. M. mali and occasionally M. fructicola
temperature, relative humidity, shade tem- produce apothecia on mummified fruit on
perature and the total daily duration of rain- the ground. All species overwinter in
fall. The model has been incorporated into infected parts of the tree. Secondary spread
an integrated apple disease warning system is by conidia produced on infected host tis-
(ADEM) and successfully used to time sue within the tree and on trees of neigh-
fungicide applications and in some cases has bouring hosts, particularly Prunus species.
resulted in improved disease control, with a Losses due to fruit decay caused by M.
40% reduction in fungicide usage (Berrie fructigena and M. fructicola increase gradu-
and Xu, 1999). ally up to harvest time and are usually asso-
ciated with injuries to the fruit. Cultivars
prone to fruit cracking, such as ‘Cox’s
18.3.3 Brown-rot diseases Orange Pippin’ and ‘James Grieve’, are
especially susceptible to infection. Infection
Several brown-rot fungi attack apple in dif- also occurs through wounds caused by
ferent parts of the world. The species and birds pecking at fruit, insects infesting fruit
their distribution are: Monilinia fructicola and hail. Fruit decay is prevented by avoid-
(Wint.) Honey in most regions except ing cultivars prone to fruit cracking, by lim-
Europe, where it is a European Union-listed iting the damage caused by birds and other
quarantine pest (Smith et al., 1992); Monilinia wounding agents and by orchard sanitation
laxa (Aderh. & Ruhl.) Honey in Asia, Europe, methods aimed at reducing the build-up of
North and South America and South Africa; inoculum.
and Monilinia fructigena (Aderh. & Ruhl.) Blossom infections from M. laxa, M. fructi-
Honey in Europe and Asia (Byrde and gena and Monilia leaf blight are controlled by
Willetts, 1977). A related species, Monilinia orchard sanitation, combined with the appli-
mali (Takahashi) Whetzel, causes Monilia leaf cation of fungicides. Blighted spurs and
blight of apple in Asia. cankers are removed and destroyed during
The brown-rot fungi cause blossom wilt, the dormant period and in the growing sea-
spur dieback, cankering and fruit rot (Plate son. Fungicides applied as the flowers begin
18.8); the incidence and severity of these to open and one or two times 5–7 days later
symptoms depend on the species of should prevent blossom blight.
18.3.4 Summer diseases has enormous potential for secondary spread

and is very difficult to control when the
Summer diseases refer to a collection of nine weather is warm and wet.
diseases that tend to be most severe from 2–3 G. cingulata also survives from year to
weeks after petal fall until harvest. They are year on dead wood and mummified apples.
most prevalent in warm and moist growing There appear to be several strains, which
regions of the world, where they can cause produce somewhat different symptoms. A
extensive losses if not controlled. They are serious leaf-spot disease, Glomerella leaf spot,
relatively minor problems in arid and cooler occurs on ‘Gala’ in Brazil, which is associ-
growing regions. ated with G. cingulata (Sutton and Sanhueza,
Of all the summer diseases, rot diseases 1998). It causes small irregular lesions
are most destructive and can cause losses of 3–12 mm in diameter on leaves and can
50% or more if not controlled. These diseases result in significant defoliation when severe.
tend to be most severe in the south-eastern It also causes small corky lesions on fruit.
USA but cause problems in other areas as This strain overwinters in leaves on the
well. The life cycles of the pathogens that orchard floor, much like apple scab, and
cause these diseases are similar, as is the infection is initiated in the late spring by air-
overall strategy for managing them. borne ascospores, which are discharged dur-
ing periods of rain. Perennation also occurs
in twig cankers and mummified apples.
18.3.4.1 Bitter rot
Glomerella leaf spot has been observed in the
Bitter rot is the most destructive of the three south-eastern USA (Gonzalez and Sutton,
rot diseases and is the most difficult of the 1999). Another strain of G. cingulata produces
three to control once an epidemic has begun. a large, firm, brown lesion on the fruit,
It is caused by Colletotrichum gloeosporioides which is not sunken, and few spores are pro-
(Penz.) & Sacc. in Penz. and Colletotrichum duced on the surface of the lesion (Shane
acutatum J.H. Simmons. These pathogens and Sutton, 1981b). It is not known whether
coexist in many orchards, whereas in others this strain causes a leaf-spot symptom.
one or the other species predominates. Management of bitter rot is based on sani-
Glomerella cingulata (Stoneman) Spauld & H. tation and a preventive spray programme. In
Shrenk is often listed as the sexual stage of C. warm, rainy, growing regions, the disease
gloeosporioides but may be a distinct taxon cannot be successfully managed without
(Shane and Sutton, 1981a; TeBeest et al., 1997). pruning to remove inoculum sources and
The Colletotrichum spp. that cause bitter facilitate drying in the tree canopy. Current-
rot survive from one growing season to season fire blight strikes need to be removed
another in mummified apples, twig cankers because they can be colonized by the bitter-
and other dead wood in the tree. In the late rot fungi and serve as an inoculum source
spring when temperatures begin to warm, late in the season. The ethylenebisdithiocar-
conidia are released, initiating infections on bamate (EBDC) fungicides are most effective,
fruit. Fruit is susceptible throughout the sea- but restrictions on their application after
son (Shane and Sutton, 1981b; Noe and petal fall in the USA have limited their use-
Starkey, 1982). Lesions begin as small, circu- fulness. Captan, ziram and thiram (applied
lar, light tan to brown spots, sometimes sur- on a 10–14-day schedule from petal fall to
rounded by a red halo. As the lesions enlarge, harvest) are the most effective fungicides
they become brown and sunken (Plate 18.9) currently registered in the USA. The benzim-
and extend into the flesh of the apple in a V- idazole fungicides are ineffective.
shaped pattern, which is a good diagnostic
technique. Fruiting structures on the surface
18.3.4.2. Black rot and bot (white) rot
of the lesion, called aecidia, produce copious
amounts of salmon-coloured conidia, which Black rot and bot rot are caused by
can be splash-dispersed to other fruit, initiat- Botryosphaeria obtusa (Schwein.) Shoemaker
ing new infections. Consequently, the disease and Botryosphaeria dothidea (Moug.) Ces. &
DeNot., respectively. Although caused by the and mushy (Plate 18.11). Under cooler tem-
same fungal genus, the diseases caused by peratures, the rot tends to be darker in
the two species are different in many ways. colour and firmer, making it more difficult
Both overwinter in dead wood, mummified to separate from black rot. Infected fruit
apples and cankers in the tree canopy and may fall or may remain attached to the tree
produce ascospores and conidia through and mummify.
most of the growing season. Both cause a Control of the black rot and bot rot is
fruit rot and cankers, but only B. obtusa based on sanitation and preventive fungicide
causes a leaf spot. Infections by B. obtusa can sprays. Colonized dead wood and mummi-
begin as early as silver tip and occur fied apples can both serve as inoculum
throughout the season, whereas infections by sources. Wood infected by the fire blight
B. dothidea occur mainly during the warm pathogen can become colonized by these
summer months. B. obtusa infections develop fungi and produce spores during the current
into firm brown lesions on the fruit and growing season. This wood should be
B. dothidea infections develop into light removed during the early summer. Prunings
brown, soft, watery rots. should be chopped by a flail mower or
B. obtusa infections can occur on the sepals pushed out of the orchard and burned.
as soon as the buds begin to open. These Fungicides should be applied every 10–14
infections spread into the fruit as they begin days from petal fall to harvest. Captan and
to mature, resulting in a firm brown rot on the benzimidazole fungicides are most effec-
the calyx end (Plate 18.10). Other fruit infec- tive; the dithiocarbamate fungicides are not
tions can occur from soon after petal fall until especially efficacious.
harvest during favourable weather (Arauz
and Sutton, 1989). Infections that occur soon
18.3.4.3 Sooty blotch and fly-speck
after petal fall first appear as small, dark pim-
ples. As lesions enlarge, they become dark Sooty blotch and fly-speck are two of the
and irregular in shape and are often sur- most common diseases in humid growing
rounded by a red halo. Eventually the entire areas. While they do not reduce yield,
fruit may become rotten and shrivel, remain- affected fruit are usually downgraded from
ing attached to the tree. Leaf spots (known as fresh-market to processing or juice grades.
frog-eye leaf spot) begin as small purple to Sooty blotch is a disease complex caused by
brown necrotic lesions, which enlarge to several fungi and is characterized by dusty
4–5 mm in diameter and are often sur- to dark colonies of fungi growing epiphyti-
rounded by a purple halo. When infections cally on the surface of the fruit (Williamson
are numerous, leaves may turn yellow and and Sutton, 2000; Plate 18.12). They range
abscise. from small discreet colonies to large, amor-
B. dothidea infections can occur from soon phous ones. Each colony has a characteristic
after petal fall until harvest (Parker and appearance, depending on the fungus
Sutton, 1993). Infections that occur early in involved. Fly-speck colonies are character-
the season often remain quiescent and do ized by numerous thyrothecia, which are
not begin developing until the soluble scattered throughout the thallus, giving it an
solids in the fruit begin increasing. This appearance of ‘fly-specks’ (Plate 18.13).
often leads growers to think that their late- Colonies range from several to many mil-
season spray programme is ineffective, limetres in diameter.
when in actuality the disease increase that Sooty blotch is a disease complex caused
they are observing is more closely related to by Peltaster fructicola Johnson, Sutton &
their spray programme earlier in the season. Hodges, Leptodontium elatius (G. Mangenot)
As lesions begin to develop on the fruit, De Hoog, Geastrumia polystigmatis Batista &
they extend in a cylindrical manner towards M.L. Farr and probably other fungi. These
the core. Once the core is invaded, the entire fungi grow superficially on the cuticle of
apple becomes infected. Rotten fruit are affected fruit. All have numerous reservoir
often light tan in colour and are very soft hosts in the wooded areas in close proximity
to orchards and most inoculum comes from for timing sprays of benzimidazole fungi-
outside the orchard. Conidia of P. fructicola cides to manage sooty blotch and fly-speck.
and G. polystigmatis are primarily water- They found that sprays of benzimidazoles
borne and are spread by wind-blown rain; could be omitted in the cover-spray pro-
conidia of L. elatius are airborne. Infection gramme until 225 h of wetting had accumu-
can occur as early as several weeks after lated. In dry years, using the model could
petal fall. Symptom expression is closely save three to five spray applications.
associated with the hours of wetting that Rosenberger (Agnello et al., 1999) and
occur in the spring. Brown and Sutton (1995) Hartman and Smigell (Hartman, 1995;
found that the first symptoms of sooty blotch Smigell and Hartman, 1998) have modified
(and fly-speck) appeared after an average of this model for their growing regions.
273 h of leaf wetting of 4 h duration or
greater accumulated following the first rain
18.3.4.4 Brooks fruit spot
which occurs 10 days after petal fall. P.
fructicola, G. polystigmatis and L. elatius all Brooks fruit spot caused by Mycosphaerella
produce conidia on fruit, which serve as pomi (Pass.) Lindau affects apples primarily
secondary inoculum. All apple cultivars are in the south-eastern and mid-Atlantic apple-
equally susceptible, but the colonies are growing regions of the USA (Sutton et al.,
more visible on light-skinned cultivars. 1987). Symptoms on fruit appear as small,
Washing and brushing in the packing line slightly sunken, superficial lesions on the
can often remove small colonies with lightly fruit (Plate 18.14). Lesions resemble cork
pigmented thalli. spot, but there is no corky tissue beneath
Fly-speck is caused by Schizothyrium pomi them. Leaf spots appear as small purple
(Mont.: Fr.) Arx (anamorph Zygophiala flecks on leaves (Plate 18.15). Affected fruit
jamaicensis E. Mason). Ascospores of S. pomi, are often downgraded from fresh-market to
which are produced in thyrothecia on reser- processing or juice grades.
voir hosts, provide the primary inoculum for M. pomi overwinters in apple leaves and
infections. While some infections occur on ascospores are discharged from pseudothe-
fruit, the most important infections occur on cia during rainfall. The ascospores are pro-
reservoir hosts, which serve as a source of duced during a distinct period, about 6
inoculum (conidia) throughout the growing weeks in duration, beginning about 2 weeks
season. Moisture and temperature require- after petal fall (Sutton et al., 1987). Symptoms
ments for colony development are similar to do not appear on fruit and leaves until 6–8
those of the fungi causing sooty blotch. While weeks after infection. There is no secondary
the fungi that cause sooty blotch grow super- spread. There are some differences among
ficially on the cuticle, Z. jamaicensis metabo- cultivars; ‘Golden Delicious’, ‘Rome Beauty’,
lizes the cuticle and the incipient thyrothecia ‘Stayman’ and ‘Idared’ are quite susceptible;
become firmly attached to it, making the ‘Delicious’ is relatively resistant. A preven-
colonies difficult to remove during washing tive fungicide program that includes a
and brushing in the packing line. There are dithiocarbamate or benzimidazole fungicide
no differences in susceptibility among culti- controls the disease.
vars to either fly-speck or sooty blotch.
Control of sooty blotch and fly-speck is
18.3.4.5 Alternaria blotch
based on sanitation to remove reservoir
hosts, pruning to open the canopy and facili- Alternaria blotch, caused by Alternaria mali
tate drying, thinning fruit clusters to Roberts (= Alternaria alternata apple patho-
improve drying, and fungicide sprays. The type), was first reported in the USA in the
benzimidazole and dithiocarbamate fungi- late 1980s (Filajdic and Sutton, 1991). The
cides are most effective; captan is not very disease affects ‘Delicious’ and cultivars with
effective. The benzimidazole fungicides have ‘Delicious’ as a parent (e.g. ‘Empire’) and is
some eradicant activity against the diseases. characterized by circular, necrotic spots on
Brown and Sutton (1995) developed a model the leaves (Plate 18.16), which, when abun-
dant, can cause extensive defoliation. lesions, often bounded by veins. Mid-shoot
Defoliation by the fungus is exacerbated by leaves are most severely affected. Lesions are
mite injury. A. mali and the European red initially pale green, over a few hours turn
mite act synergistically to increase defolia- chocolate brown and, as they age, often
tion (Filajdic et al., 1995). When defoliation is appear tan (Plate 18.18). Severely affected
extensive, fruit size and soluble solids are leaves turn yellow in a few days and abscise.
reduced. Fruit symptoms are usually limited The disorder appears following periods of
to small, corky, dark lesions, often associated cloudy, cool weather during the summer and
with the lenticels. can result in 50% or more defoliation. It has
The fungus overwinters in leaves on the been associated with an increase in the pro-
orchard floor and to a lesser extent in leaf duction of gibberellins, which is triggered by
buds. Infection by airborne conidia occurs environmental factors. Necrotic leaf blotch
within a month of petal fall if temperature can be controlled by applications of heavy
and moisture conditions are favourable. The metal-containing fungicides, such as ziram,
numerous secondary cycles that can occur thiram, mancozeb and Bordeaux, or foliar
throughout the growing season may result in nutrient sprays (e.g. zinc oxide).
extensive defoliation.
Control of the disease is based primarily
on preventive control of mites to minimize 18.3.5 Phytophthora crown and root rot
defoliation. Most fungicides currently regis-
tered for apples in the USA, with the excep- Phytophthora crown and root rot is a poten-
tion of the strobilurins, have little effect on tially serious soil-borne disease wherever
Alternaria blotch. apples are grown. The disease is prevalent
when susceptible rootstocks are planted in
heavy soils or in areas of poor soil drainage.
18.3.4.6 Black pox
Infected trees exhibit a variety of symptoms.
Black pox, caused by Helminthosporium papu- During summer, foliage on infected trees
losum Berg., is a problem, especially on may appear light green. As the season pro-
‘Golden Delicious’ in the south-eastern USA. gresses, leaves on infected trees turn a red-
It is characterized by small, black, slightly dish-bronze. Branches on infected trees
sunken lesions, 3–9 mm in diameter on the typically have stunted leaves and poor ter-
fruit (Plate 18.17). Leaf spots begin as red minal growth. Fruit on infected trees is
haloes with light green centres, enlarge to smaller than normal and colours prema-
1.5–11.0 mm and turn tan to brown with pur- turely. Accurate diagnosis of Phytophthora
ple borders (Taylor, 1963). H. papulosum over- crown and root rot frequently requires dig-
winters in twig lesions. Conidia produced in ging in order to expose the upper portions
these lesions initiate infections during the of the root system. Infected tissue is reddish
summer. Black pox is one of the least-studied brown and delineated from healthy tissue
summer diseases, and conditions favouring by a definite margin (Plate 18.19). Diseased
infection are not known. Preventive fungicide trees are often randomly distributed
sprays of dithiocarbamate or benzimidazole throughout a planting. The disease is some-
fungicides provide effective control. times confused with the rootstock phase of
fire blight.
The Phytophthora species present, soil
18.3.4.7 Necrotic leaf blotch of ‘Golden
moisture and temperature, relative resistance
Delicious’
of the rootstock and contamination of nursery
Necrotic leaf blotch is a physiological disor- stock all affect the incidence and severity of
der that affects primarily ‘Golden Delicious’ the disease (Welsh, 1942; Sewell and Wilson,
and its progeny (Sutton and Sanhueza, 1998) 1959; Browne, 1984; Jeffers and Aldwinckle,
and is a particular problem on the new culti- 1986; Ogawa and English, 1991). The disease
var ‘Pacific Rose’. It is characterized by the can result from infection by any of several
sudden appearance of large, irregular species in the genus Phytophthora: P. cactorum,
P. cryptogea, P. cambivora, P. megasperma, P. 18.3.6 European or Nectria canker

syringae, P. citricola and P. drechsleri. These
fungi can persist in soil for extended periods Nectria or European canker is circumglobal
of time as thick-walled sexual spores, called in distribution and particularly problematic
oospores. When soil becomes saturated and in northern Europe and parts of South
temperatures are conducive for sporulation, America (Grove, 1990). The disease is consid-
oospores germinate to produce sporangia. ered minor in most of North America, but
Within sporangia are numerous unicellular can be a serious problem in the production
motile spores that are liberated into the soil areas of coastal California. The disease is
water. The spores, known as zoospores, swim characterized by zonate cankers located on
short distances in the soil pore spaces or can the main trunk or scaffold limbs (Plate
be transported longer distances by runoff or 18.20). Young infected tissue is darker than
irrigation water. When zoospores contact surrounding healthy tissue. Limb or tree
roots of susceptible hosts, they germinate and death can occur from girdling, which results
establish new infections. Zoospores are liber- from canker enlargement. A layer of callus
ated only when soil is saturated. Prolonged tissue is formed annually around each
flooding favours the production of sporangia canker. The fungus invades callus tissue and
and dissemination of zoospores and may also spreads to healthy tissue outside the callus
predispose the host to infection. The majority layer. Over a period of years this results in
of new infections are initiated between the cankers with characteristic zonate appear-
pink stage of blossom development and the ances. In some areas nurseries are important
beginning of shoot elongation. The optimum sources of infected trees.
temperature for disease development The disease is caused by Nectria galligena
depends upon the Phytophthora species pre- Bres. The fungus produces ascospores and
sent, but in general soil temperatures between conidia in orange-red fruiting-bodies pro-
20 and 30°C favour the disease. duced in twig, branch or trunk cankers. Spores
Phytophthora crown and root rot is man- are produced in a gelatinous matrix and dis-
aged through the integration of chemical and persed by the impaction of water droplets.
cultural practices. Two of the most important The disease can be aggravated by over-the-
decisions of the producer are to plant canopy irrigation. Infection occurs through
pathogen-free nursery stock and to avoid the leaf scars and pruning wounds. Nectria canker
use of susceptible rootstocks. Rootstocks is favoured by cool, moist weather.
vary in their susceptibility to the various An integrated approach is required for
species of Phytophthora. The vast majority of successful disease management. Diseased
rootstock-susceptibility evaluations have wood should be removed from the orchard
been conducted using P. cactorum. The and destroyed. Young infected trees should be
MM.104 and MM.106 rootstocks are particu- severely pruned or removed. In some areas
larly susceptible to the disease, while M.9 is diseased bark is removed to prevent spore
relatively resistant. During planting, orchard production. Copper fungicides (e.g. Bordeaux
managers should ensure that the graft union mixture) are sometimes applied prior to
is not in contact with soil. Heavy or poorly autumn rains in order to protect leaf scars.
drained soils should be avoided. In irrigated
areas, water should be managed to avoid
prolonged flooding and the presence of 18.4 Postharvest Diseases
standing water around the base of trees.
Plantings are sometimes established on Apples are also host to a multitude of
raised beds in order to facilitate drainage postharvest diseases and disorders. The for-
around the base of trees. Acylalanine and mer are caused exclusively by pathogenic
ethyl phosphonate fungicides available for fungi. Two of the more serious postharvest
control of this disease are applied as soil diseases are blue mould and grey mould,
drenches or foliar sprays, respectively which are the first and second most impor-
(Jeffers and Wilcox, 1990). tant postharvest diseases of apple, respec-
tively. Most postharvest pathogens invade ties or severe tree decline and even death.
fruit wounded during harvest, shipping or The degree of affliction depends greatly on
handling. Therefore, the management of the rootstock and scion selection, pathogen
postharvest diseases begins with very careful isolate and climate. Viruses or virus-like
fruit harvest, bin sanitation and transport. agents that incite severe symptoms on some
Fruit bins used for transport should be free apple selections will cause no obvious symp-
of soil, leaves and rotten fruit debris. Fruit toms on others. However, even in the latter
should be picked individually and placed case, significant yield reductions have been
very gently into bins. Orchard access roads documented in the few studies undertaken
should be smooth and free of ruts and major to study such effects (van Oosten, 1983).
bumps. Bins should be transported at speeds Thus, infections by viruses and virus-like
that will not result in shifting or bouncing. agents are important to the commercial pro-
Additional means of postharvest disease duction of apples.
management include packing-house sanita- Although most of these agents do not
tion, rapid immediate postharvest cooling, spread naturally, many are common in com-
fungicide drenches and the provision of ade- mercial orchards. The universal presence of
quate tree nutrition. many of these agents is due to collecting
grafting material from infected but appar-
ently symptomless trees while preparing for
18.4.1 Blue mould vegetative propagation of new trees. The
propagation of trees from infected sources
Blue mould, which is caused by fungi in the ultimately affects yield and, under certain
genus Penicillium, is the most common environmental circumstances or clone com-
postharvest disease of apple (Rosenberger, binations, results in orchards that are not
1990; Plate 18.21). Decayed flesh is soft and economically productive due to fruit-quality
watery and separates readily from healthy issues or tree decline. The only reasonable
tissue. Infected epidermal tissue is light to method for controlling these diseases is the
dark brown. Infected fruit are malodorous. initial use of virus-tested propagation
The causal organism produces blue or blue- materials.
green conidia on the surface of infected fruit. Quick and reliable diagnostic assays are
Penicillium spp. are present in orchard soils available for a few of the viruses and virus-
and on decayed fruit in the orchard and like agents that infect apple. However, for
packing-house and in postharvest drench the most part, detection of such infections is
solutions and flume water. Fruit infection a cumbersome and laborious proposition.
typically occurs through wounds. Wounding Furthermore, only a few of the agents that
should be prevented during harvest, ship- incite these diseases have been identified.
ping and processing. Infested fruit bins and The use of modern laboratory diagnosis is
warehouses should be disinfested before precluded until such identifications are
processing fruit. made. Most assays for these agents are con-
ducted by inoculative assays on sensitive
woody indicator selections; these tests
18.5 Diseases Caused by Viruses, require 2 months to 3 years to complete.
Viroids, Phytoplasmas and Other Virus- Aetiological studies of viruses that infect
like Agents woody plants are greatly enhanced if the
virus is first transmitted to a herbaceous
Viruses or virus-like agents incite over 50 host. Unfortunately, many of the virus-like
described diseases of apple. These agents agents that elicit disease in apple trees have
include viruses, viroids and phytoplasmas not been successfully transmitted to herba-
and several graft-transmissible agents that ceous plants and therefore are called ‘non-
have yet to be identified. Some of these dis- sap-transmissible’. This may simply mean
eases have great economic consequences in that the correct combinations of host plants
that they are associated with fruit deformi- and transfer conditions have not yet been
found. The significance of this limitation is share many important qualities, we may be
that progress towards characterization of able to predict the behaviour of that
these pathogens has been dramatically pathogen based on the epidemiology of a
slowed. Nevertheless, the non-sap-transmis- few well-studied pathogens in the same
sible viruses of fruit trees present a very group.
interesting and challenging group of viruses, The list of apple diseases caused by
since relatively little is known about them. ‘virus-like’ agents is alarming in its length.
Contemporary methods in molecular biol- However, many of these disease names were
ogy have accelerated investigation of these applied locally to a particular symptom and
pathogens. thus many of the names are duplicative.
Pathogens are frequently referred to as Also, many of these diseases have very lim-
viruses based on the ability to transmit the ited distribution. Only diseases that remain
pathogen by grafting and/or budding and of a more general importance will be dis-
the absence of any other obvious pathogens. cussed in detail. For an expanded list of
This simplistic distinction is often incorrect. reported diseases of apple that are induced
Many disease-causing agents are more cor- by virus-like agents, see Németh (1986).
rectly referred to as ‘virus-like’ agents since
no direct association between a pathogen
and disease has been established – that is, 18.5.1 Chlorotic leaf spot
Koch’s postulates have not been fulfilled.
The disease names commonly used in the Relatively few well-characterized viruses
literature are descriptive of the disease and are routinely found in the Malus of com-
host but provide little information about the merce (Table 18.3). Each of these viruses
pathogen. Consequently, a pathogen or vari- incites disease with major characteristics on
ant thereof may be associated with different selected host cultivars that help identify the
disease names, depending on the symptom disease agent.
produced, the latter frequently affected by The causal agent apple chlorotic leaf-
cultivar and environment. With advances in spot trichovirus (ACLSV) is one of the most
our ability to isolate and characterize fruit- widely distributed viruses of fruit trees.
tree pathogens at the molecular level, this Although first described as a virus in
complicated structure of nomenclature is apples, it was subsequently found to be
evolving to reflect more accurately the very widespread in stone fruits, where it
nature of the pathogen, rather than the can cause severe losses in production.
symptoms that they elicit. This is of practical Although the interaction of ACLSV with
as well as academic importance. By relating many commercial apple cultivars is latent, it
a pathogen to other members of a group that was first discovered associated with the
Table 18.3. Diseases of apple with known virus aetiology and their vectors.
Known means of
Disease Causal agent transmission
Apple chlorotic leaf spot Apple chlorotic leaf-spot trichovirus Grafting

Apple decline Apple stem-grooving capillovirus Grafting
(on Virginia crab)
Apple mosaic Apple mosaic ilarvirus Grafting
Apple mosaic (Tulare) Tulare apple-mosaic ilarvirus Grafting
Apple stem grooving Apple stem-grooving capillovirus Grafting
Apple stem pitting Apple stem-pitting foveavirus Grafting
Apple union necrosis Tomato ringspot nepovirus Nematodes and grafting
Flat apple Cherry rasp-leaf nepovirus Nematodes and grafting
Spy decline Apple stem pitting foveavirus Grafting
decline and death of new Malus breeding cultivars, but appears on rootstocks with
lines in an apple-scab-resistance breeding crab apple in their heritage.
programme. Over the past few decades, ASGV can be diagnosed by herbaceous
there have been several examples where the indexing on C. quinoa, although this method
interaction of the virus with specific root- is not very reliable. Woody indexing on
stocks is not latent. The top-working dis- Malus micromalus GMAL273.a or M. sylvestris
ease of Japan is believed to be a cv. ‘Virginia Crab’ (Howell et al., 1995) or
hypersensitive reaction of rootstocks (Malus detection by RT-PCR is the preferred method
prunifolia var. ringo) to isolates of ACLSV of testing (Kummert et al., 1995; Kinard et al.,
found in contaminated apple scions from 1996).
North America. This resulted in significant
failure of mature trees (reviewed in Mink,
1989). The death of cells at the graft union 18.5.3 Flat apple
results in weakened trees with reduced
vigour and productivity and which are very This disease is caused by cherry rasp-leaf
susceptible to wind damage. nepovirus (CRLV) (Parish, 1977), a virus
ACLSV is diagnosed by serological test- that is believed to be native to western
ing, indexing on herbaceous hosts North America, ranging from Utah to
(Chenopodium quinoa) or budding to Malus southern British Columbia. CRLV is
indicator species, usually Malus sylvestris cv. believed to have originated in one or two
R12740-7A (also known as ‘Russian’). In the species of native vegetation and subse-
latter case, chlorotic leaf symptoms appear quently been transmitted into horticultur-
soon after new leaves develop unless the ally important crop plants, such as apple
ambient temperature rises above 20°C. Low and cherry. The symptoms in apple are
virus titre and inhibitors in Malus often limit most striking on the fruit of ‘Red Delicious’
the use of serology or indexing on herba- and related cultivars. The length of the fruit
ceous plants. Molecular techniques of diag- is significantly reduced to produce a fruit
nosis, such as RT-PCR, are becoming more that is squat, and the stem cavity is almost
common methods of diagnosis (Kummert et absent (Plate 18.22). At first, only a few fruit
al., 1995; Kinard et al., 1996). The extreme will be affected, but eventually the entire
strain variation of ACLSV has limited the tree will be involved.
widespread acceptance of serological and CRLV is transmitted by the nematode
molecular methods. Xiphinema americanum (sensu lato) (Wagnon et
al., 1968). Therefore, secondary spread of the
disease is relatively slow. Apple trees
18.5.2 Apple decline (on ‘Virginia Crab’) planted on sites previously planted to rasp-
leaf-diseased cherry trees can become
Infection of most apple trees with apple infected. In addition to fruit trees, many
stem-grooving capillovirus (ASGV) is latent orchard weeds, such as dandelions and plan-
– that is, there are no acute symptoms visi- tains, are hosts, although they exhibit no
ble. However, when infected scion material symptoms of the disease. This makes elimi-
is grafted on to sensitive rootstocks, such as nation of the virus from an infected orchard
M. sylvestris cv. ‘Virginia Crab’, growth and very difficult. The alternative hosts and/or
development of the ‘Virginia Crab’ wood the nematode vectors would have to be elim-
cylinder is impaired and deep grooves inated to protect replanted trees.
appear under the bark. The vascular tissue Since CRLV achieves high concentration
of the ‘Virginia Crab’ often becomes in young shoots, it can be readily detected
necrotic, resulting in weakness and a visible by serology and by mechanical inoculation
brown line at the graft union. The trees of C. quinoa with extracts from young leaves
often break at this necrotic union (Németh, in early spring. Leaf symptoms and tip
1986). Stem grooving caused by ASGV is necrosis on C. quinoa usually develop 3 days
not seen on current commercial apple after inoculation.
18.5.4 Apple mosaic used to detect ASPV. Epinasty and decline

may be observed after 1 year when ‘Spy227’ is
Apple mosaic ilarvirus (ApMV) induces dra- the recipient. ‘Radiant’ crab apple is a more
matic white bands or patterns on leaves of reliable detection host. It is sensitive to a wide
many apple cultivars (Plate 18.23). Affected range of ASPV isolates and displays severe
leaves may have irregular distribution epinasty and decline within 6 weeks under
through the tree or even on individual limbs. controlled greenhouse conditions. The devel-
The severity of the symptoms can vary dra- opment of stem-pitting symptoms on
matically from year to year, with symptoms ‘Virginia Crab’ is also reliable, but at least two
being almost undetectable in seasons with growing seasons are required for trustworthy
high temperatures during the growing sea- results. Fruit with ridges and flutes are pro-
son. This disease can result in significant duced if the ‘Virginia Crab’ is allowed to bear.
production losses (ranging up to 40%) Molecular methods are also available for the
depending on the cultivar, virus isolate and detection of ASPV. RT-PCR (Nemchinov et al.,
environment (Posnette and Cropley, 1956). 1998) is sensitive and offers faster diagnosis
ApMV is detected by serological assays relative to woody indexing.
but results (when apple tissue is tested) are
variable. Therefore, serological testing is best
limited to confirming the identity of the 18.5.6 Union necrosis
pathogen in herbaceous indexing. ApMV can
also be detected by inoculating field-grown Union necrosis is caused by tomato ringspot
trees of ‘Golden Delicious’ apple. Herbaceous nepovirus (TmRSV) (Stouffer and Uyemoto,
indexing on C. quinoa is possible but quite 1976). Symptoms of infection generally do
unreliable. Molecular methods of detection, not appear until the tree reaches bearing age,
such as RT-PCR (Rowhani et al., 1995), are at which time the tree assumes an unthrifty
becoming more widely accepted, as the growth habit. The leaves and bark are red-
methods are evaluated with more and varied dish in colour, and bloom and fruit set are
isolates of the virus. abnormally high. Depending on the root-
stock/scion combination, the results of infec-
tion may range from very mild to a rapid
18.5.5 Stem pitting decline in tree health, resulting in death. In
severe reactions (‘Red Delicious’ on MM.106),
The viral nature of apple stem pitting was the union of rootstock and scion will reveal a
discovered soon after the Second World War dark necrotic line spanning the union, with
in mid-western North America (Guengerich soft, spongy, orange bark flanking the union.
and Millikan, 1956). Production of crab This disease is common in several locales
apples had become uneconomic and old crab along the east coast of North America, and
apple rootstocks were budded over to stan- has been reported sporadically in the Pacific
dard cultivars, such as ‘Delicious’. Two or Northwest of the USA. TmRSV is transmitted
three years after budding, the crab apple by dagger nematodes, X. americanum (sensu
rootstocks became pitted and severely weak- lato), which are abundant in eastern fruit-
ened. This is an example where the ‘latent’ growing areas of North America (Stouffer
virus in the ‘Delicious’ scion resulted in sig- and Powell, 1989). Other species belonging to
nificant economic damage when the variety this nematode group, Xiphinema revesi and
was budded on to rootstocks with crab apple Xiphinema californicum, are also known to be
in their heritage. Apple stem-pitting fovea- vectors of TmRSV in orchards.
virus (ASPV), the causal agent of apple stem- TmRSV is easily transmitted from young
pitting disease, is thought to have been leaves to C. quinoa by mechanical inoculation
distributed worldwide in contaminated with crude extracts. Prunus tomentosa is a diag-
apple and pear propagation material. nostic woody host for this virus. Alternatively,
Woody indexing on any one of a variety of serological and RT-PCR (Griesbach, 1995)
Malus species is the method most frequently assays can be used to detect it.
18.6 Diseases Caused by Phytoplasmas 18.6.2 Chat fruit

(Table 18.4)
Apple chat fruit is believed to be caused by
18.6.1 Proliferation phytoplasma infection. Sensitive cultivars
develop undersized fruit, many of which
Apple proliferation occurs throughout much drop in June. The remaining fruit do not
of Europe, but has not been reported in the develop colour properly and may exhibit
western hemisphere. The disease is caused dark water-soaked spots. Many of the cur-
by an apple proliferation phytoplasma. The rent commercial cultivars do not express any
pattern of occurrence in European orchards symptoms of apple chat fruit. The pathogen
suggests that the pathogen is harboured in is detected by budding on to ‘Lord
the native weed population, from which it Lambourne’. An incubation period of up to 3
moves into the orchards. The vector of this years is required for reliable detection. Until
pathogen is at least one species of leafhop- a firm association of a biological agent with
per, Fieberiella florii. This disease reduces fruit apple chat-fruit disease can be established,
size by up to 30–70% and fruit have longer the use of biological indicators remains the
than normal peduncles. However, the most only means of detection.
striking symptom, for which the disease is
named, is the proliferation of vegetative
shoots, otherwise known as witches’-broom.
18.7 Diseases of Apple Caused by Viroids
In addition, apple leaves exhibit greatly
(Table 18.5)
enlarged stipules. Because the disease is
debilitating and insect-vectored, apple prolif-
Viroids are sub-virus particles that are diffi-
eration disease has important quarantine sig-
cult to detect. Since proteins are not part of
nificance in the western hemisphere.
their structure, serological methods are not
The most widely accepted detection
method for apple proliferation phyto- useful. Woody indexing and molecular
plasma has been woody indexing in the detection methods are commonly used to
field. Bark patches from roots of the test diagnose these pathogens.
plant are budded on to ‘Golden Delicious’
and the tree is observed for 2 years for the
appearance of witches’-broom and the 18.7.1 Blister bark
enlarged stipules. However, the long obser-
vation period and concern about reliable Apple blister bark is characterized by areas
transmission of the pathogen encouraged of the bark taking on the appearance of
the development of alternative detection orange tissue-paper. The disease can be
methods. Detection by PCR (Lorenz et al., induced by apple fruit-crinkle viroid
1995) is becoming widely accepted and the (AFCVd) (Ito et al., 1993). As the underlying
techniques are becoming increasingly tissue becomes desiccated, the bark cracks
refined to improve the reliability of molecu- and peels. Scarring of the underlying tissue
lar methods (Carraro et al., 1998; occurs. Several mineral deficiencies can
Skrzeczkowski et al., 2001). PCR is now the mimic infection by AFCVd, so proper diag-
preferred test method. nosis is important.
Table 18.4. Diseases of apple suspected to be caused by phytoplasma.
Known means of
Apple chat fruit Phytoplasma Grafting

Apple proliferation Apple-proliferation phytoplasma Leafhoppers and grafting
Table 18.5. Diseases of apple known to be caused by viroids.
Known means of
Apple blister bark Apple fruit-crinkle viroid (AFCVd) Grafting

Apple dimple fruit Apple dimple-fruit viroid (ADFVd) Grafting
Apple fruit crinkle Apple fruit-crinkle viroid (AFCVd) Grafting
Apple scar skin Apple scar-skin viroid (ASSVd) Grafting
Dapple apple Apple scar-skin viroid (ASSVd) Grafting
18.7.2 Dapple apple than fruit from uninfected trees. Diagnosis is

performed as described for dapple-apple dis-
Dapple apple aptly describes one of the disease. The dapple and scar-skin diseases
eases caused by apple scar-skin viroid (ASSVd). evoked by ASSVd appear to be dependent
Affected fruit, especially near the calyx end, upon climatic and cultivar differences. The
bear small circular spots, which enlarge and original isolates of each produce similar dis-
increasingly contrast with the background eases on fruiting trees of ‘Scarlet Pimpernel’
colour as the fruit matures (Plate 18.24). Larger (W.E. Howell, Washington State University,
spots may coalesce to produce dappling or a unpublished data).
broad zone of discoloration. There are no pro-
nounced leaf or bark symptoms. Since many
commercial cultivars do not express symptoms, 18.7.3 Dimple fruit
this disease has been particularly problematic
when old orchards are top-worked to change Apple dimple fruit caused by apple dimple-
cultivars. The original cultivar may not express fruit viroid (ADFVd) has been observed in
symptoms, whereas the new cultivar does. Pear several commercial cultivars of southern
also appears to be a symptomless carrier of Italy (DiSerio et al., 1996). The disease
ASSVd; the role of pear as a symptomless car- induces depressions on the fruit surface as it
rier of this pathogen in apple-disease aetiology matures. Under the depression will be an
is unknown. ‘Stark’s Earliest’ (‘Scarlet area of necrotic tissue.
Pimpernel’) or ‘Delicious’ can be used for
woody indexing, and three crops must be
observed for accurate assessment of viroid sta- 18.7.4 Fruit crinkle
tus. ‘Scarlet Pimpernel’ grown at 18ºC under
constant light for 2 months will yield diagnostic Apple fruit-crinkle disease is caused by iso-
symptoms on its foliage (Skrzeczkowski et al., lates of AFCVd (Ito et al., 1993). The viroid
1993). Molecular techniques are preferred for was described in Japan, where fruit symp-
their speed and reliability. Both RT-PCR toms are most severe on the cultivar ‘Ohrin’.
(Hadidi and Yang, 1990) and hybridization This pathogen can be detected by woody
(Hadidi et al., 1991) assays are used for ASSVd. indexing on the apple cultivars ‘Delicious’
Apple scar-skin disease is a more severe or, preferably, ‘NY5822’, where it induces
disease caused by ASSVd, as compared with symptoms of blister bark (Ito et al., 1993).
the milder one referred to as dapple apple.
Symptoms usually begin at the distal portion
of the fruit and the severity increases as the 18.8 ’Virus-like’ or Graft-transmissible
fruit matures. Small discoloured circles merge Diseases of Apple with No Known Causal
to form large green-brown to brown patches Agents (Table 18.6)
(Plate 18.25). Eventually, the brown patches
become necrotic and fissures appear on the There are many diseases of apple for which
fruit. Diseased fruit are significantly smaller no pathogen has been identified. The disease
Table 18.6. Graft-transmissible diseases of apple for which there are no known causal agents.a
Apple blister bark Apple green crinkle Apple ringspot

Apple dead spur Apple horseshoe wound Apple rosette
Apple decline Apple internal bark necrosis Apple rough skin
Apple false sting Apple leaf pucker Apple rubbery wood
Apple flat limb Apple ring russet Apple pustule canker
Apple freckle scurf Apple ring-line pattern
aThere are many more described but rare graft-transmissible diseases of apple (Németh, 1986).
names are generally descriptive of the symp- may appear on only one or two limbs of an
toms. Some may be caused by pathogens infected tree, and there are no acute foliar
described above but, until more information symptoms associated with the disease
is obtained about their aetiology, the causal (Thomsen, 1989). Budding and grafting of
agents remain an enigma. Since the agents for contaminated propagation material is the
these diseases have not been identified, con- major mechanism by which apple green-
firmation of the pathogen depends on symp- crinkle disease is spread. If field spread
tom expression on susceptible cultivars. All occurs, it is very slow and possibly by root
are graft-transmissible. Furthermore, as most grafting only.
of these diseases are spread primarily
through the use of infected propagation
material, the creation of virus-certification 18.8.2 Rubbery wood
programmes over the past 40 years has essen-
tially eliminated many of these diseases from Apple rubbery wood affects 2-year-old wood
commercial apple production. Still a few con- of apple and pear trees. On sensitive culti-
tinue to cause concern. vars, branches develop that are very pliable
and droop under their own weights
(Waterworth and Fridlund, 1989). Affected
18.8.1 Green crinkle limbs are also very sensitive to cold and frost
damage. The disease originated in Europe
Apple green-crinkle disease, a severely and was later introduced to North America
debilitating disease of some apple cultivars, with infected rootstock. Many older root-
is always associated with trees infected with stocks introduced from Europe were uni-
ASGV, ASPV and ACLSV. The individual formly infected with rubbery wood. In
viruses have not been separated to deter- countries with active certification pro-
mine if this disease symptom is induced by a grammes, this disease has become increas-
mixture of viruses, by a particular isolate of ingly rare. The degree of symptom
one of the viruses or by another as yet development is dependent on climate. The
uncharacterized pathogen. Apple green-crin- pliable limbs and poor growth are more
kle disease is diagnosed by woody indexing extreme in moderate climates, relative to
on the apple indicator ‘Golden Delicious’. warmer environments. No obvious symp-
The trees are observed for the development toms are associated with many commercial
of fruit symptoms during the following three cultivars. Nevertheless, fruit yield and qual-
crops. Symptoms begin to appear after the ity can be adversely affected (van Oosten,
fruit reaches 1–2 cm in diameter. Fruits 1983). The pathogen has been assumed to be
develop deep depressions and distortions, a phytoplasma, based on electron-
which become more severe as the fruits microscopic examination of affected tissues,
mature. Cracks may develop in pits and but attempts to confirm this association by
crevices (Plate 18.26). Depressions are linked molecular techniques have been contradic-
through the flesh to the vascular system by tory (Poggi Pollini et al., 1995; Bertaccini et
discoloured tissue. Severe fruit symptoms al., 1998). The disease is detected by indexing
on ‘Lord Lambourne’, followed by 3 years of 18.9 Control Measures

observation for the development of limbs
lacking normal lignification. Very few viruses or virus-like agents of apple
Apple flat limb is believed to be caused disease spread naturally in the orchard. This
by the same agent that causes apple rubbery means that the most efficient and cost-effec-
wood (see above). Apple flat limb is only tive control measure for apple virus diseases
observed in those areas where sensitive culti- is the use of propagation material and plants
vars, such as ‘Gravenstein’, are grown. The that are certified to be free from viruses. Once
disease results in flattening of the shoots and the trees are planted in the orchard, they
branches on limbs that are 2–3 years old and should, for the most part, remain virus-free.
eventually leads to deep furrows that Still, some spread of these pathogens occurs
become more severe as the branches become in orchards via tree-to-tree root grafts. Such
older (Fridlund and Waterworth, 1989). In root grafts provide effective means by which
addition to reduced vigour and production, pathogens can move from an infected tree to
the weak limbs are susceptible to easy break- the next. However, this process is relatively
ing. Severely affected limbs are also more slow and involves spread only to adjacent
susceptible to winter or frost damage. The trees. Thus, if a diseased tree is detected in an
disease is detected by woody indexing on established orchard, it may be prudent to
‘Gravenstein’ or ‘Lord Lambourne’, followed remove adjacent trees to eliminate the virus.
by 3 years of observation. In those cases where some form of natural
field transmission exists, diseases can be
much more difficult to control. Again, initial
planting of certified virus-free material is the
18.8.3 Dead spur
first step in maintaining a healthy orchard.
In European countries, where apple prolifer-
Apple dead spur was originally observed in
ation phytoplasma is prevalent, regular
the western USA, but has since been
insecticide applications to control the
reported throughout North America and in
leafhopper vector can significantly reduce
Europe and Asia (Parish, 1989). The charac- the incidence of this disease (Kunze, 1976).
teristic symptom is death of fruiting spurs, The two diseases of apple whose causal
with the resulting development of long seg- agents are known to be transmitted by nema-
ments of blind wood. This is most pro- todes, flat apple and union necrosis, can be
nounced on the interior of the tree, so the very difficult to control. Exclusion of the virus
canopy in the centre of the tree is sparse, initially is crucial since, once the virus is pre-
with fruiting spurs at the shoot tips. The dis- sent, it is very difficult to eliminate it from an
ease is spread through the use of infected orchard. Dagger nematodes, Xiphinema spp.,
propagation material. No natural spread has are nearly universal and difficult to eradicate.
been observed. The only means by which the Fumigation and soil pretreatment before plant-
diagnosis based on symptoms can be con- ing will reduce but not eliminate these nema-
firmed is bud-inoculating spur-type todes from soil. Leaving the ground planted in
‘Delicious’ trees. Symptoms will develop in plants that are not virus hosts for 1–2 years
the third or fourth year after inoculation. before replanting will help prevent transmis-
Since this disease is difficult to diagnose, it sion of these viruses to the young trees. The
may be overlooked in virus-testing pro- nematodes lose the ability to transmit these
grammes. Like green-crinkle disease, dead viruses with each moult. Since many weed
spur has not been observed in the absence of plants common in the orchard floor, such as
ACLSV, ASPV or ASGV. Therefore, although dandelion and plantain, act as reservoirs of
the dead-spur agent poses a concern for the these two nepoviruses, intense weed control is
safe propagation of healthy apple trees, elimi- required during this period of fallow. If these
nation from propagation programmes of trees viruses and their nematode vectors are present
with these three ‘sentinel’ viruses may well in an area, it is best to select rootstock/scion
indicate freedom from this and other graft- combinations that offer protection against the
transmitted diseases of unknown aetiology. nematode and/or virus disease.
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19 Ecology and Management of Apple

Arthropod Pests
Elizabeth H. Beers,1 D. Max Suckling,2 Ronald J. Prokopy3 and

Jesús Avilla4
1Washington State University, Tree Fruit Research and Extension Center, Wenatchee,
Washington, USA; 2The Horticulture and Food Research Institute of New Zealand Ltd,
Canterbury, New Zealand; 3Department of Entomology, University of Massachusetts,
Amherst, Massachusetts, USA; 4Centro UdL-IRTA de R+D de Lleida, Universidad de
Lleida, Lleida, Spain

19.2 Systems of Pest Management 490
19.2.1 Pesticide-based 490
19.2.2 Integrated pest management 499
19.3 Fruit Feeders 501
19.3.1 Direct pests of buds and fruitlets 502
19.3.2 Mature-fruit feeders 503
19.4 Foliage Feeders 509
19.4.1 Mesophyll stylet feeders 510
19.4.2 Bulk leaf feeders 512
19.5 Structural Feeders 512
19.5.1 Superficial woody-tissue and shoot feeders 512
19.5.2 Wood-boring insects 513
19.5.3 Root-system pests 514
19.6 Conclusion 514
19.1 Introduction ered pests at some point in time. This sur-

vey referred to one orchard in a temperate
Apples present a distinct challenge to inte- production zone in central North America,
grated pest management (IPM), due in part and we can only presume the total for the
to their perennial growth habit and physical world is far greater. Despite this, only a
complexity. The various organs of the tree’s dozen or so arthropods in any given region
structure provide multiple habitats suitable are considered serious or chronic pests. A
for arthropod colonization. In one study few, such as the codling moth, the European
(Oatman et al., 1964), 763 species of arthro- red mite and, to a lesser extent, the two-
pods were discovered using apple as a host spotted spider mite are pests virtually
plant. While many of these were transitory, wherever apples are grown; others are
perhaps 100 or so species have been consid- strictly regional pests.
490 E.H. Beers et al.
When the pest complexes are viewed as The classification of pests in this chapter
a whole, a pattern of ecological homo- is necessarily an arbitrary choice. We refer
logues emerges. These homologues may be to arthropod taxa, but, for pest-manage-
closely related species, or unrelated taxa ment purposes, the taxon is not necessarily
that have similar feeding habits. The the most useful unit. Our approach has
tetranychid mite complex in the Pacific been more crop-centred, in that groupings
north-west (Tetranychus urticae Koch, have been made on the basis of damage
Panonychus ulmi (Koch) and Tetranychus type (Fig. 19.1), which is in turn usually
mcdanieli McGregor) all feed in the same highly related to its potential economic
manner and cause a similar type of foliar importance. Within some of the larger
damage (Beers et al., 1993). The leaf-roller groups (fruit feeders), we have grouped
complex (moths in the family Tortricidae) pests by time of attack or by type of damage
all feed on leaves and the surface of apple caused. Overarching the crop and produc-
fruits. Weevils (e.g. the plum curculio tivity issues, we have superimposed the
Conotrachelus nenuphar (Herbst)) and thrips ecological niche and ecological homologue
(the western flower thrips, Frankliniella occi- concepts in an attempt to make the
dentalis (Pergande)) are examples of two plant–herbivore relationship clearer.
unrelated taxa that cause similar types of
damage (surface feeding and oviposition,
leaving a superficial scar) and at about the 19.2 Systems of Pest Management
same period in fruit development (during
or shortly after bloom). 19.2.1 Pesticide-based
A number of pest species are strictly
monophagous on apple (e.g. Aphis pomi De The discovery and commercialization of
Geer), while others are oligophagous or synthetic organic pesticides in the latter half
even highly polyphagous (e.g. T. urticae). of the 20th century represented a major
The degree of host specialization does not qualitative change in pest management. For
appear to be related to pest status. One of the first time since the beginning of agricul-
the key pests worldwide (codling moth, ture, producers had a broad range of highly
Cydia pomonella (L.)) is moderately effective and relatively inexpensive prod-
oligophagous, feeding primarily on a few ucts to use for insect control (Table 19.1).
species of Rosaceae and one member (wal- Their ease of use and often long residual
nut) of the Juglandaceae. However, many toxicity to pests made them very popular
species exhibit a certain degree of plasticity and, to some extent, the applications were
in their feeding behaviour and are capable an insurance policy against pest damage.
of shifting hosts or expanding their host The euphoria was short-lived, as resistance
range over time. An example is the apple problems began developing, sometimes
maggot, Rhagoletis pomonella Walsh, in west- within a few seasons’ use. The organochlo-
ern North America. A host shift was recently rines, introduced to agriculture after the
demonstrated for this species (from apple to Second World War, were largely supplanted
cherry) (Jones et al., 1989), even though a by the organophosphates, carbamates and
closely related species, Rhagoletis indifferens pyrethroids within a few decades. The
Curran, already occupied this niche in this problems associated with the use of these
region (Utah). Apple is an introduced crop products became apparent after a relatively
in the majority of the areas where it is short time, including environmental persis-
grown, so the pest complex of any given tence and damage (especially the
region is typically a mixture of pests from organochlorines), mammalian toxicity (e.g.
the native region that have been introduced applicator and farm-worker safety, espe-
over time (many before strict quarantine cially the organophosphates), possible con-
regulations were imposed) and native pests sumer effects from residues on foods
that have adapted to using apple as a host (carcinogenicity, teratogenicity, mutagenic-
(e.g. apple maggot). ity or chronic neural effects), and destruc-
Apple Arthropod Pests 491
Fig. 19.1. Examples of arthropod pests attacking various parts of the tree. Clockwise from top: scale (feed
on bark); aphids (phloem feeders in shoots and leaves); leafhoppers (pierce mesophyll cells and remove
contents); woolly apple aphid galls (on roots); bark beetles (attack trunk and major scaffolds); leaf-rollers
(feed on fruit surface and leaves); codling moth (feeds internally in fruit); plum curculio (oviposits and scars
young fruitlets). (Illustration by G. Steffan.)
tion of pests’ natural enemies and selection tive tactics was minimal, because of the effi-
for resistant pest populations. There were cacy of the new pesticides. Non-pesticidal
clear economic benefits driving the use of tactics with some degree of promise were
these materials: 30–50% damage from dismissed because of their relatively higher
codling moth in the latter part of the lead expense, lower efficacy or greater complexity
arsenate era (1940s) was common (Driggers, of implementation. The concept of mating
1937), whereas the economic threshold for disruption, well established by the 1970s
this pest today is generally set at < 1%. (Roelofs, 1979), was not registered for use on
Despite this, the disenchantment with these apples in North America until the early 1990s,
materials has been growing steadily since and is still not registered in some European
the 1950s. countries. Similarly, the sterile-insect tech-
One of the side-effects of the pesticide- nique, although demonstrated as feasible for
based era was that the bulk of entomological codling-moth control in the 1960s (Proverbs
research was directed at the development et al., 1966), was not implemented in tree
and optimum use of the new pesticides, and fruit on a large commercial scale until the
basic biology and biological-control research early 1990s, and then only on a limited
slowed considerably. The search for alterna- acreage in British Columbia, Canada.
Apples - Chap 19
492
Table 19.1. Historical use of insecticides and acaricides in apple.
Type
(I = insecticide, Use period
Class/pesticidea A = acaricide) (approximate) Comments
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Inorganic
Lead Arsenate I 1890s–1950s Once the sole control measure for codling moth and other pests, this compound
was used for > 50 years until resistance occurred and replacement insecticides
11:01 am
became available. Soil residues are still present
Sulphur I/A Late 1800s–present Often applied with lime as a safener, this material is still widely used for both
day arthropod pests and diseases. Used in late winter or early spring, it can be
phytotoxic
Cryolite I Used briefly during periods of codling-moth resistance; occasional use in organic
Page 492
production
Dinitro Compounds Several compounds in this group have been used, but DNOC was the most common
E.H. Beers et al.

Dinitro-o-cresol (DNOC) I/A 1930s–1970s Highly phytotoxic; thus use was confined to dormant sprays. Used with oil to
control aphid eggs and overwintering scale. No longer permitted in Europe (2000)
DN-111 A 1940s–early 1950s A summer acaricide. Phytotoxic
Botanicals As a group, these were once the primary pesticides allowed in organic produc-
tion; some are being withdrawn (see individual chemicals). Widely variable in
terms of mammalian toxicity
Neem I 1980s–present Derived from the seeds of neem (tree) (Azadirachta indica A. Juss); has
antifeedant, repellency and/or growth-regulator influence on many orders of
insects
Ryania extracts (main I 1950s–present Ground bark of a tropical shrub (Ryania spp.); once widely used for codling-moth
active ingredient ryanodine) control, it still has a limited place in organic apple production
Rotenone I/A 1930s–present A neurotoxin best known for its toxicity to fish; no longer allowed in most organic
certification programmes. Component of roots of tropical plants (e.g. Derris spp.).
Controls a wide range of arthropod pests
Pyrethins I/A Ancient times Several derivatives of flowers in the genus Chrysanthemum. Control of a wide range
to present of insects and mites. Residues disappear very quickly. Little commercial orchard
use except organic
Nicotine I A highly poisonous substance derived from Nicotiana spp., used commonly in the
early part of the century for aphid and other soft-bodied insect control. Usually as
nicotine sulphate
Apples - Chap 19
Chlorinated hydrocarbons Generally a very persistent group of neurotoxic compounds in soil, water and animal
tissues; very few still in use today for this reason. Mammalian toxicity relatively low
DDT
(dichlorodiphenyltrichloroethane) I Mid-1940s–1970s The most recognizable name in this class, subject of the book Silent Spring.
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Highly persistent, originally with a very broad spectrum of activity. Primary target
was codling moth, but it created severe mite flare-ups. Toxic to many beneficial
insects
TDE (DDD) I Physical and chemical properties similar to DDT; more effective than DDT against
11:01 am
(dichlorodiphenyldichloroethane) red-banded leaf-roller
Benzene hexachloride (BHC) I Used primarily pre-bloom for aphid control; in season use could give fruit an off
flavour
Lindane I 1940s–present Purer gamma isomer of BHC, more widely used
Methoxychlor I 1940s–present Use in tree fruit currently limited to leaf-miner control (as a premix with malathion)
Page 493
Endrin I 1960s Limited use against Lepidoptera
Endosulfan I/A 1950s–present One of the few remaining chlorinated hydrocarbon compounds still widely used
Apple Arthropod Pests

(primarily in USA). Control of sucking, chewing, boring insects; some acaricidal
activity, especially rust mites
Dicofol A 1950s–present Related to DDT. Highly toxic to phytoseiid mites, less used currently
DMC (dichloromethylbenzhydrol) A 1950s Related to DDT. Limited availability and high cost
Ethyldichlorobenzilate A 1950s Limited use due to phytotoxicity
Organophosphates Broad-spectrum neurotoxins introduced after Second World War, many members
acutely toxic to mammals. Many were acaricidal when first used, but resistance
developed after a few seasons’ use. Once the most prevalent group used on tree
fruits, they are gradually being replaced by new compounds
TEPP (tetraethylpyrophosphate) I/A Very highly toxic to mammals, but short-lived residues. Used against aphids,
mites, scales and Lepidoptera
Azinphosmethyl I 1950s–present Broad-spectrum and widely used for 30–40 years; fairly high mammalian toxicity;
uses currently being restricted
Diazinon I 1950s–present Broad-spectrum, moderate mammalian toxicity; also available to home-owners
Malathion I 1950s–present One of the lowest mammalian-toxicity compounds in this group; little used in
commercial production any more. Short residual, thus preharvest use is popular
Chlorpyrifos-ethyl I 1960–present Widely used for pre-bloom aphid control and post-bloom Lepidoptera control
Chlorpyrifos-methyl I 1960s–present Control of various foliar pests (lepidopterous, aphids, scales)
Ethion I 1950s–1980s Some use pre-bloom for scale and aphids
Carbophenothion I/(A) 1950s Somewhat phytotoxic, more limited spectrum of activity than azinphosmethyl and
parathion
493
Continued
Apples - Chap 19
494
Type
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Methidathion I 1950s–present Minor use pre-bloom against San Jose scale (USA)
Ethyl parathion I/(A) Late 1940s–1990s Highly toxic, broad-spectrum and once widely used, this material was withdrawn
from the market in some countries in the 1990s. Originally also acaricidal
11:01 am
Methyl parathion I 1940s–1990s Similar in toxicity and spectrum to ethyl parathion; often sold as an encapsulated
formulation to prolong the residue; withdrawn from the market in the 1990s
Phosmet I Early 1970s–present Moderately broad activity, similar to azinphosmethyl, but lower worker hazard
Demeton I/(A) Mid-1950s–late 1980s A systemic material used primarily for aphid control. Originally acaricidal
Phorate I/(A) 1950s Systemic in both foliar and soil applications. Originally acaricidal. Potentially
Page 494
phytotoxic
Phosphamidon I/(A) 1950s–1980s Systemic. Widely used as an aphicide, originally acaricidal. Marginally phytotoxic
Mevinphos I Mid-1950s–mid- Systemic. Extreme acute oral and dermal toxicity to mammals, but short residual.
E.H. Beers et al.

1990s Used primarily as an aphicide, some Lepidoptera activity
Dimethoate I/A Late 1960s–present Systemic. Used for aphid and Lygus control
Phosalone I/A 1960s–present Broad-spectrum pesticide. Control of Lepidoptera and Diptera
Carbamates Neurotoxins with a slightly different mode of activity from that of the organophos-
phates
Carbaryl I/A 1950s– present Broad-spectrum insecticide, low mammalian toxicity; widely used as a fruit thinner
as well as an insecticide. Some eriophyid activity, toxic to phytoseiids, causing
spider-mite outbreaks
Methomyl I 1970s–present Much higher mammalian toxicity, used primarily for control of Lepidoptera
Oxamyl I/A Mid-1980s–present A more toxic carbamate, sometimes used for Lepidoptera; also toxic to both
phytophagous and predatory mites
Formetanate hydrochloride I/A 1970s–present Effective against mites, thrips, some Hemiptera/Homoptera and Lepidoptera;
toxic to predatory mites
Pirimicarb I 1970s–present Selective systemic insecticide used primarily as an aphicide (except USA). Low
toxicity to natural enemies
Organotins Acaricides widely used in the 1970s and 1980s; resistance problems curtailed use
Azocyclotin A 1970s–present Long-acting acaricide with contact action
Cyhexatin (hexakis) A 1970s–mid-1980s Widely used until resistance became widespread; withdrawn from the US market
in 1987
Fenbutatin oxide A 1970s–present Similar in activity to cyhexatin, apparent cross-resistance to that compound
Apples - Chap 19
Pyrethroids A group based on the activity of natural pyrethins, but more active and with longer
residual. Generally-broad spectrum insecticides/acaricidies with low mammalian
toxicity, but famous for acute toxicity to phytoseiid mites. Useful near to harvest
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due to their small safe-to-harvest interval (few days)
Fenvalerate/esfenvalerate I/A 1970s–present Targeted Lepidoptera, but other pests also controlled (Hemiptera/Homoptera).
Esfenvalerate was a more active isomer of fenvalerate, replacing it in the 1980s
Permethrin I 1970s–present Control of fruit- and leaf-eating Lepidoptera and Coleoptera
11:01 am
Acrinathrin A/I 1990s–present Mainly used as an acaricide against European red mite. Good insecticidal activity
against thrips
Bifenthrin I/A 1980s–present Mainly used as an insecticide against Lepidoptera
Deltamethrin I 1970s–present Broad-spectrum insecticide, also used against fruit flies
Flucythrinate I 1980s–present Mainly against Lepidoptera and Homoptera
Page 495
Lambda-cyhalothrin I 1980s–present Broad-spectrum
Tau-fluvalinate I/A 1980s–present It replaced fluvalinate. Mainly used against lepidopterous and aphid pests

Microbial insecticides Materials that produce disease in the insect host; very specific, thus mammalian
toxicity is low
Bacillus thuringiensis (Bt) I 1980s–present Bacteria that produce an exotoxin, which, when ingested, causes gut paralysis.
subsp. kurstaki Specific to lepidopterous larvae. Primarily for leaf-rollers
Codling moth granulovirus I 1980s–present Viral disease specific to codling moth; used primarily in Europe as a ‘soft’
(CpGV) insecticide supplement to codling-moth control. Low persistence
Adoxophyes orana I 1990s–present Viral disease specific to summer fruit tortrix larvae. More effective against first-
granulovirus (AoGV) instar larvae
Beauvaria bassiana I 1980s–present Fungal disease; dependent on weather conditions; little commercial use as yet
Macrocyclic lactones A relatively new group of compounds whose active ingredient is from toxins pro-
duced by soil microorganisms; large complex molecules, some are semi-synthetic
Abamectin I/A 1980s–present Derived from Streptomyces avermitilis; controls mites, leaf-miners, some areas
report control of leafhopper
Spinosad I 1990s–present Derived from Saccharopolyspora spinosa; leaf-roller and leaf-miner control, also
thrips and possibly some tephritid fruit flies
Milbemectin I/A Derived from Streptomyces hygroscopicus. Activity spectrum similar to that of
abamectin; not yet registered in USA/Europe
Polynactins A Derived from Streptomyces aureus. Control of spider mites
Continued
495
Apples - Chap 19
496
Type
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Insect growth regulators A newer group of insecticides attacking various points in the insect’s hormonal
system, thus making them specific to invertebrates and largely non-toxic to mam-
mals. Targets are mostly Lepidoptera, some Homoptera
Benzoylureas (diflubenzuron, I/A 1970s–present These compounds act as chitin-synthesis inhibitors. Used mainly against leaf-
11:01 am
hexaflumuron, flufenoxuron, and fruit-eating lepidopterous larvae (codling moth, leaf-rollers and leaf-miners);
triflumuron, lufenuron, some have some effect against rust mites (lufenuron) or spider mites
teflubenzuron) (flufenoxuron); resistance to diflubenzuron has been reported in Europe; never
registered in the USA
Fenoxycarb I 1980s–present Although chemically a carbamate, it acts as a juvenile hormone analogue, with a
Page 496
strong juvenile hormone-like activity, inhibiting metamorphosis to the adult stage
and interfering with the moulting of early-instar larvae; widely used in Europe from
E.H. Beers et al.

the 1980s against codling moth and leaf-rollers; never registered in the USA
Tebufenozide I 1990s–present Ecdysone agonist,b which acts by binding to the ecdysone receptor protein. As a
consequence, the moulting process is lethally accelerated. Used in Europe for the
control of codling moth and leaf-rollers
Methoxyfenozide I 1990s–present Ecdysone agonist; more active than tebufenozide; codling moth, leaf-rollers, leaf-
miners
Pyriproxyfen I 2000–present Juvenile hormone analogue, good scale and other Homoptera activity, some
suppression of Lepidoptera
Nicotinoids Neurotoxins that act at the nicotinyl site; a newer group of insecticides, fairly
broad activity spectrum
Imidacloprid I 1980s–present The earliest registration of the group; widely used for aphid control; also effective
against other Homoptera, including leafhoppers and mealybugs. Also toxic to
apple maggot
Thiamethoxam I 2001–present Recently registered; activity spectrum includes Lepidoptera and
Hemiptera/Homoptera
Chlorinated sulphur acaricides
Aramite A 1950s A chlorinated sulphite
Chlorfenson (Ovex) A 1950s A chlorinated sulphonate, somewhat phytototoxic
Genite 923, Mitox, Fenson A 1950s–1960s Closely related chlorinated sulphur compounds
Sulphenone A 1950s A chlorinated sulphone. Phytotoxic
Tetradifon A 1960s–present Long residual effect, translaminar activity. Also ovicidal
Apples - Chap 19
Miscellaneous synthetic organic pesticides
Oxythioquinox I/A 1960s–present A heterocyclic carbonate, used primarily as an acaricide, but with some activity
against psylla and mildew
Indoxacarb I 2001–present A carbamate-like compound, primarily used against Lepidoptera
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Pyridaben A/I 1990s–present Used mainly in apple orchards as an acaricide against European red mite. It is an
inhibitor of the electron transport at mitochondrial level (METIc). High knock-down
effect and long residual activity to all mobile stages. Toxic to phytoseiids. Risk of
developing resistance
11:01 am
Tebufenpyrad A 1990s–present Another METI acaricide, but with some activity against summer eggs and also a
translaminar action. Toxic to phytoseiids. Risk of developing resistance
Fenazaquin A 1990s–present Another METI acaricide with some activity against summer eggs. Toxic to
phytoseiids. Risk of developing resistance
Fenpyroximate A 1990s–present Acaricide active against Tetranychidae and some effect against Eriophyidae. It
Page 497
acts as a growth regulator. Moderately toxic to phytoseiids
Chlordimeform A 1970s A chlorinated phenamidine

Propargite A 1970s–present A relatively selective acaricide, withdrawn from the US market in the 1990s due to
worker dermatitis problems. Still in use in Europe
Hexythiazox A 1990s–present It has ovicidal, larvicidal and nymphicidal activity, and also sterilizes females;
highly selective, but potential for resistance found soon after introduction. It
inhibits the synthesis of chitin
Clofentezine A 1980s–present Primarily ovicidal (it inhibits the development of the embryo) and some action
against newly hatched larvae, long persistent and highly selective, but potential
for resistance found soon after introduction
Amitraz A/I 1970s–present Mainly used as an acaricide, to control all stages of tetranychid and eriophyid mites
Other Materials of this type, some of which have been used for over a century, are enjoying
a resurgence of interest, due to their low environmental and human health impact
Oil (petroleum) I/A 1880s–present Highly refined narrow-cut petroleum products with emulsifiers added; broad
activity against soft-bodied insects and some repellent activity (especially
oviposition). Often used as an adjuvant
Oil (plant-derived) Early 1900s Mainly used as stickers for other pesticides
Oil (animal-derived) Early 1900s–present Primarily fish-oil. Widely used against codling moth in the early part of the
century, used in organic production today to some extent
Kaolin clay I/A Late 1990s–present Also known as particle film technology (PFT), this recently introduced compound
has a broad spectrum of activity. Probably repellent, or masks plant host
Diatomaceous earth I 1950s–present An abrasive silica-containing material mined from deposits of skeletons of marine
microorganisms; many industrial uses; little used in apple production
497
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Type
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Soap I/A 1950s–present Fatty acid derivatives that are broadly toxic to soft-bodied insects; not widely used
because of short residual, high cost and potential phytotoxicity. Phosphate-based
E.H. Beers et al.

laundry soaps are also insecticidal
Mating disruption I 1990s–present Synthetic chemicals mimicking natural insect pheromones. Not direct toxicants,
but reduce insect populations; registered as ‘pesticides’. Available for codling
moth, oriental fruit moth and some leaf-rollers
Mass trapping I 1990s–present Use of pheromones (or other attractants) to catch a high percentage of the adult
population. Available for some wood-borers and tephritid flies
Attract and kill I 1990s–present Use of pheromones (or other attractants) to attract adults to a droplet of sticky
material that contains a rapid knock-down insecticide. Available for codling moth
aPrimary reference material from western USA and Europe. Key references include Tomlin (2000), ACTA (2001), De Liñán (2001), selected chapters in Fisher
and Upshall, (1976). See References.
bAlso known as a moult-accelerating compound (MAC).
cMETI, mitochondrial electron transport inhibitor.
19.2.2 Integrated pest management from the orchard and the value of the crop. The
second is that of sampling pest populations in
The problems with the so-called ‘pesticide order to arrive at some numerical or risk-based
treadmill’ were part of the impetus to reassessment of the population. With these ele-
examine and reorganize pest-management ments in place, a comparison is made between
efforts. The use of pesticides engendered a the cost of some control measure and the pro-
pest-by-pest approach, with little regard for jected value of crop loss from insect damage.
the effect of the sprays on the rest of the The point at which the two are equal is called
agroecosystem (let alone the environment or the economic injury level (EIL) (Stern et al.,
consumer). With the realization that these dis- 1959). As a general principle, the producer
junctive efforts were in some cases working wants to ensure that the insect population
against each other, a framework of thought does not exceed the EIL (because preventable
was developed to try simultaneously to economic loss occurs), nor is there any particu-
account for multiple effects and to solve mul- lar benefit in merely breaking even. Ideally,
tiple problems. This theoretical framework the producer needs to forecast the future
became known as ‘integrated pest manage- insect population from the current one (based
ment’, or IPM (Stern et al., 1959). Although on previous experience or population growth
there are a number of variants (Steiner et al., models) and, when it is clear that the EIL will
1977), this is still the predominant philosophy be exceeded at some future date, the control
governing apple pest research. measure is the preferred course of action.
The guiding philosophy behind IPM was Not surprisingly, all elements of this sys-
the optimization and harmonization of tactics tem are fraught with uncertainty. Unless the
to achieve ‘the best economic, environmental, producer is growing his/her fruit under con-
and social’ outcome (Rabb, 1972). While this tract, the future value of the fruit is unknown.
seems straightforward enough, turning this Indeed, for decisions made early in the sea-
philosophy into practice has been an ongoing son, even the size of the crop is uncertain.
and occasionally hotly disputed process. One Insect population growth is influenced by
overriding difficulty has been evaluating the many factors, including the action of natural
relative value of a practice where there are enemies, the influence of weather conditions,
clear economic consequences (usually for the sprays aimed at other pests or diseases and
producer) and more nebulous, but potentially the tree vigour. All of these modify the
far-reaching, consequences for society at insect’s innate ability to reproduce (the intrin-
large. Where the quality and quantity of food sic rate of increase (Birch, 1948)), which is the
production overall are an overriding issue, interaction of the number of progeny per
the value of less expensive or more abundant female, the time to first reproduction and the
food has often outweighed the more long- sex ratio. Examples of accurate models that
term environmental and social issues. are actually in use in apple production are
However, in affluent countries with ample few, if any; however, there is usually a good
food supply and relative economic wealth, sense of the potential growth factor of an
the conflict becomes more acute. This is gen- insect population from one generation to the
erally reflected in the increasing interest in next in the absence of control measures. In
IPM, integrated fruit production (IFP) and several cases (e.g. tetranychid mites and
organic production in Europe, the Americas gracillariid leaf-miners), the modifying effect
and parts of Asia. of natural enemies is partially quantified,
A number of key concepts of IPM form the such that, at a given predator : pest ratio
foundation of most apple pest-management (Croft, 1975; Avilla et al., 1993) or percentage
programmes (Metcalf and Luckmann, 1975). parasitism (Beers et al., 1993), a reasonable
The first fundamental concept is to develop estimate of whether the population will
some quantitative relationship between the require treatment can be made.
pest population and the loss in yield or pro- Another aspect to sampling insect popu-
ductivity. This loss must then be assigned an lations is monitoring their phenological
economic value, based on the projected yield development in order to determine the opti-
mum timing for control measures. Insects 19.2.2.1 IPM tactics

are poikilotherms and speed or slow their
development in response to ambient tem- In one sense, almost any pest-control tactic
perature. This principle underlies the con- may potentially have a place in an IPM
cept of physiological time, using some type framework. Some tactics tend to be more
of time–temperature summation (e.g. often associated with IPM or viewed more
degree-days). Computer simulations of favourably. It should be noted at the outset
development (degree-day models) were that the use of insecticides and acaricides, in
developed for many pest species (e.g. the appropriate circumstances, is considered
codling moth (Fig. 19.2)). This degree-day a legitimate IPM tactic. Increasingly, IPM is
model has been applied most often to the defining the characteristics of appropriate
determination of optimum timing of pesti- pesticides more and more narrowly. In any
cide applications, but is equally applicable case, pesticide use must always be context-
to (for example) distributing mating-disrup- sensitive: the insect must have reached some
tion dispensers in the orchard or releasing a critical population level to warrant treat-
biocontrol agent. Since monitoring some ment; the optimum timing and placement of
species may be difficult (due to extremely the material must have been considered; the
low population levels) or time-consuming, most appropriate compound must be chosen
phenological models have been developed in light of its effects on natural enemies and
to facilitate the process. These models, often other pests in the orchard. In addition, fac-
driven by fairly simple temperature inputs tors such as worker and environmental
(daily maxima and minima) and some ini- safety are being given more weight in the
tialization point (often the first capture of an decision-making process.
adult in a pheromone or visual/odour trap), Biological control is considered in many
provide producers with greatly improved ways to be the ideal pest-management tac-
accuracy of determining insect-stage devel- tic, because it tends to be environmentally
opment. They do not, however, tell the pro- innocuous, self-sustaining and low cost.
ducer anything about the need for control Each of these characteristics may depend a
measures, which must be accomplished by great deal on the system in question. The
other means. low environmental impact of biological
100
90
% Development (codling moth)
80
70
% Male flight
60
% Egg hatch
50
40
30
20
10
0
0 500 1000 1500 2000 2500
Accumulated degree-days
Fig.19.2. Codling-moth degree-day model. Degrees are calculated using a horizontal cut-off sine-wave
method, with lower and upper temperatures of 10 and 31°C, respectively (Brunner et al., 1982).
control was formerly considered dogma; such as aphids. Cultivars or strains that have
recent studies (Follett and Duan, 2000), reduced terminal growth, such as the spur-
however, point out that ecosystem disrup- type cultivars, may play the same role. In
tion from imported organisms can be exten- general, however, producers prioritize plant
sive and unexpected. growth and productivity in their orchard-
Conservation biological control is management practices, which may conflict
arguably the easiest and thus most fre- with the optimal pest-control practice.
quently pursued. This approach uses a nat- Host-plant resistance, while frequently
ural enemy species that already occurs in the used in field crops, has played a very small
region and makes the environment more role in arthropod-pest management of
favourable for its growth and development. orchard crops. The horticultural characteris-
This can include cultivating plants in the tics, especially precocity, productivity,
vicinity of the orchard that provide an alter- flavour and storability, are the primary dri-
native insect host or habitat or avoiding pes- vers of cultivar choice. One notable excep-
ticides that are toxic to one or more life tion is the use of resistant rootstocks for
stages. The latter is often referred to as ‘inte- woolly apple aphid.
grated control’ or the integration of biologi- Ultimately, IPM can be viewed as just
cal and chemical control tactics. Classical another evolutionary step in our overall
biological control is the importation of a nat- problem-solving process in agriculture. More
ural enemy, often from the region where the recently, theories have emerged (primarily in
crop originates, which has the capacity to Europe and New Zealand) that take the next
provide complete economic control of the logical step of integration to the entire pro-
pest in question. The purest form of this type duction system – integrated fruit production,
is in minimally managed systems, where or IFP (Boller et al., 1998; see Chapter 21). To
pesticide use for other pests does not disrupt an extent, this may be viewed as a reincarna-
the imported natural enemy. Examples of tion of the organic-production philosophy
this type are rare in tree fruits, because the (see Chapter 22), which also encompasses all
use of at least some pesticides is ubiquitous. aspects of the production system but with
However, there is still an interest in the the additional caveat of restricting the mate-
importation of natural enemies, which, if rials used to only naturally occurring, mini-
established, become candidates for conserva- mally processed products (in terms of
tion biological control. The last methods pesticides, plant-growth regulators and fer-
involve ongoing releases of artificially reared tilizers).
natural enemies; these can occur either occa-
sionally (augmentative) or in the form of a
‘biological pesticide’ (inundative). Because 19.3 Fruit Feeders
the expense of rearing natural enemies can
be considerable, the latter two methods have This group of insects attacks the fruit
been little implemented. directly, leaving either feeding scars or deep
Cultural control involves manipulating entries, potentially serving as an infection
the orchard or the immediate environment to site for pathogens. The EILs for these pests
reduce pest numbers or mitigate pest dam- are relatively straightforward for fresh-mar-
age. Irrigation may reduce water stress and ket fruit, because virtually all defects are
allow arthropod-stressed trees to produce removed during packing. The issue is some-
better than they could otherwise. Orchard- what clouded for processing fruit, where
floor management (e.g. the mix of plants in some level of damage, especially healed sur-
the row middles) may allow more natural face damage, does not detract from the util-
enemies to build up in the cover crop and be ity or quality of the fruit. Overall,
available to reduce arboreal pest popula- pest-management programmes have
tions. Reducing fertilization so that vegeta- focused most intensely on this group of
tive growth is minimized may slow the pests because of their clear and apparent
population growth of flush-feeding insects, effect on usable yield.
The fruit may be attacked at almost any C. nenuphar (Herbst), which has become a
point during the growing season, from early key pest of apple and other pome and stone
in the bud stage to harvest. Fruit attacked fruit in its native range of eastern and mid-
early in the season is more likely to abscise western North America. In addition, several
naturally, or it can be selectively thinned species of Rynchites are local or sporadic
during hand-thinning. Fruit attacked during pests in Europe.
the mid-season is more likely to stay on the Apple-blossom weevil adults feed on
tree and thus has a higher likelihood of being developing apple buds in spring. Feeding is
harvested. Fruit attacked very late may gen- followed by oviposition and larval feeding
erate sufficient ethylene to abscise prema- on the bases of flower petals, resulting in
turely and has a slightly reduced chance of sterility and a brown-capped appearance of
entering the packing or processing plant. the flowers. Low to moderate populations
Clearly, excessive amounts of fruit drop just may act as natural blossom thinners. Large
before harvest will have a detrimental effect populations, more common in recent years,
on yield. can overthin the crop. Plum-curculio adults
likewise feed on developing apple buds in
spring but also feed upon and then oviposit
19.3.1 Direct pests of buds and fruitlets into young fruitlets, where larvae tunnel and
cause most injured fruitlets to drop. Injured
19.3.1.1 Noctuids (Lepidoptera: Noctuidae) fruit remaining on trees are scarred by the
feeding and ovipositional wounds, which
There is a complex of species in this family in usually render injured fruit unmarketable.
which the young larvae feed on developing Whereas apple-blossom weevils and north-
buds and fruitlets. The feeding damage can ern populations of plum curculios have one
prevent development, cause premature generation per year, more southern popula-
abscission or leave deep scars that distort the tions of plum curculio have an additional
fruit. This group, called the green fruit- generation and threaten not only fruitlets but
worms in North America, include Orthosia also apples approaching maturity.
hibisci (Guenée), Amphipyra pyrimadoides An understanding of the ecology of these
(Guenée) and Lithophane antennata (Walker). weevil species is the key to successful man-
Several species, such as Orthosia incerta agement (Vincent et al., 1999). Both species
(Hufnagel), may be found in Europe, can build into large populations on unman-
depending on the region (Carter, 1984). aged host trees. In some locales, plum cur-
These pests may be regionally important, but culio annually infests 90% of the fruit on
are generally considered minor. Pheromones unmanaged trees. Although resident verte-
may be used to monitor their flight to help brate and invertebrate predators, parasitoids
predict phenology and relative abundance, and pathogens do have some impact, the
e.g. Graphania mutans in New Zealand degree of population suppression by these
(Burnip et al., 1995). biocontrol agents has generally been insuffi-
cient to maintain infestations below levels
that threaten the quality of buds or fruitlets.
19.3.1.2 Weevils (Coleoptera: Curculionidae,
Fortunately, adults of both species have
Attelabidae)
rather limited flight capability, usually no
Although several different species of weevils more than a few hundred metres. Even so,
are known to feed on buds, fruit, foliage and many blocks of apple trees in Europe and
woody tissue of apple trees, only two are eastern and midwestern North America
considered to be major pests against which have at least one border exposed to suffi-
apple growers take specific action. These are cient numbers of nearby unmanaged hosts
the apple blossom weevil, Anthonomus pomo- to constitute high susceptibility to invasion.
rum (L.), a native and widespread pest of Another important ecological consideration
apples (and occasionally pears) in Europe is overwintering, which occurs in the adult
(Toepfer et al., 1999), and the plum curculio, stage, when individuals move in autumn
from infested trees to protected sites beneath 1976). With the exception of L. lineolaris,
fallen leaves, bark or debris at margins of most of the apple-feeding mirids are facul-
nearby woods or hedgerows. Finally, when tatively predacious and thus are considered
overwintered adults migrate into orchards natural enemies as well as pests.
in spring, there is a strong propensity for
establishment on perimeter trees and succes-
19.3.1.4 Thrips (Thysanoptera: Thripidae)
sively less propensity for movement on to
interior trees with increasing distance from Thrips are serious and widespread crop
the perimeter. pests worldwide, but have few representa-
Application of organophosphate or other tives in the apple pest complex. The most
insecticides timed to coincide with pulses of common species is F. occidentalis (Pergande).
adult immigration continues to be the main The adults are attracted to blooming plants
approach to managing both of these pests. and are often present in the orchard on
Because there still exists no truly effective blooming weeds. When apple blossoms
trap for monitoring immigrant adults open, they move to developing fruits. Their
(Prokopy et al., 1999), timing of application is feeding activities (sucking mouth-parts)
based on degree-day models that predict cause a condition called ‘pansy spot’ on sen-
periods of immigration (Reissig et al., 1998). sitive cultivars, and they leave a small ovipo-
Improved understanding of the ecology of sition scar in the centre of the pansy. The
these species has facilitated excellent damage is most apparent on light-coloured
orchard-wide control using a much-reduced cultivars, often colouring over on deeply
amount of material through restricting appli- coloured sports (Plate 19.2). The pear thrips,
cation to only those orchard trees most likely Taeniothrips inconsequens (Uzel) is primarily a
to become infested, i.e. trees within 20 m or pest of pear and sugar-maple, but is an occa-
less of the perimeter (Vincent et al., 1997). sional pest of apple.
19.3.1.3 Mirids (Hemiptera: Miridae) 19.3.1.5 Sawflies (Hymenoptera:

Like the weevils, the serious mirid pests of Tenthredinidae)
apple are orchard invaders, completing the Hymenopterous pests of apple are few in
majority of their life cycle outside the number (see also late-season direct fruit
orchard and immigrating only during brief feeders). Hoplocampa testudinea (Klug), the
periods to feed on fruit. This presents an apple sawfly, is a widespread and sometimes
additional challenge to pest management in serious pest of apple in Europe (Giraud et al.,
that the grower is forced to respond reac- 1996), although elevated natural mortalities
tively, rather than being able to take proac- may be caused by various fungi and the ich-
tive steps in management. The tarnished neumonid Lathrolestes marginatus (Jaworska,
plant bug or Lygus bug (Lygus lineolaris 1992). The adults appear during bloom and
Palisot de Beauvois) (Plate 19.1) is a spo- lay eggs in the flower, giving rise to larvae
radic pest of apple. It pierces the develop-
that burrow and feed in the fruit. The adults
ing fruitlet with its piercing–sucking
may be monitored with white sticky panels.
mouth-parts, leaving a deep, inverted dim-
ple on the mature fruit. Although the
mullein plant bug (Campylomma verbasci
19.3.2 Mature-fruit feeders
(Meyer)) feeds in a similar way, it leaves a
raised corky wart on the fruit. Several other
19.3.2.1 Codling moth
pests in the same group occur in different
areas of Europe and North America, includ- BIOLOGY Codling moth, C. pomonella (L.), is
ing the genera Lygocoris, Lygidea, the main direct pest of apples worldwide
Heterocordylus (Boivin and Stewart, 1982), and has been extensively studied (e.g.
Campyloneura, Plesiocoris, Blepharidopterus http://ippc.orst.edu/codlingmoth) (Plate
(Alford, 1984) and Atractotomus (MacPhee, 19.3). It is not reported as present in Japan,
Taiwan, Korea or eastern China, but is other- walnut host plants (Barnes, 1991). The
wise cosmopolitan. It is present in the urban removal of alternative or abandoned host
areas of the Brazilian apple-growing area, trees can therefore make an important contri-
but it has not yet invaded the orchards. bution to control by reducing migration of
There are typically between one and four the pest into smaller orchards.
generations per year, depending on the cli-
mate. The level of infestation on untreated DETECTION AND INSPECTION METHODS Pheromone
apple trees can reach 100% of fruit infested, traps have been used for detecting adult male
with evidence of multiple ‘stings’ or larval codling moths since the initial pheromone
attacks. The economic threshold for codling identification (Roelofs et al., 1971). This is one
moth is low (c. 1% damaged fruit), even for of the best understood and most widely used
crops that are not exported. These factors pheromone monitoring systems. A number of
have combined to make this pest one of the different management approaches have been
greatest scourges for apple growers. It is also based on pheromone-trap detection of males,
one of the most researched and consequently including forecasting female moth flight and
best understood insect pests. The absence of oviposition from sustained male flight activity,
the insect from Asian growing regions has used with day-degree accumulation (Riedl,
led to stringent procedures, including fumi- 1976), spray thresholds based on the number
gation of apples and other potential host of moths in standard traps (Wearing and
fruit with methyl bromide (e.g. Maindonald Charles, 1978) and the use of traps to deter-
et al., 1992). mine the efficacy of mating disruption, some-
Female moths oviposit single eggs on or times with lures with higher pheromone loads
near developing fruit. Larvae hatch out and to overcome the pheromone background
locate apples on the basis of an apple fruit (Charmillot, 1990).
volatile, (E,E)-α-farnesene (Sutherland and Cardboard bands applied at the right time
Hutchins, 1972). Larvae then begin to enter around tree trunks to collect diapausing lar-
the fruit and make their way to the core to vae are useful for estimating the number of
feed on the seeds, like other members of the larvae per tree (Eyer, 1937) and have been
genus Cydia (Witzgall et al., 1996b). The widely used in research and by organic
entrance hole is frequently plugged with growers for cultural control. They may be
frass. Mature larvae emerge from the fruit especially useful for comparing the larval
with a characteristic exit hole. Diapausing populations from year to year in a given
fifth-instar larvae overwinter in cocoons in orchard. Direct observation of damaged
suitably protected locations under the bark of apples during the growing season is another
the host tree or on the ground. Factors con- obvious method of monitoring the pest pop-
tributing to population regulation of codling ulation, although detection of a direct pest at
moth have been the subject of considerable harvest is usually too late for economic pro-
research. There appears to be general accep- duction where there is a single generation.
tance of the findings of Geier (1963) that lim-
ited supply of fruit and overwintering sites CHEMICAL CONTROL For much of the 20th cen-
are the key factors limiting codling-moth tury, chemical control was the most wide-
populations on unmanaged trees. spread method of pest control. However,
The main recorded hosts are apple, after usage and selection for populations
European pear, nashi (Asian pear), Chinese with genetic resistance to arsenic (Hough,
pear and quince. Walnut and plum are con- 1928), followed by the same pattern with
sistently attacked, while peach, nectarine dichlorodiphenyltrichloroethene (DDT)
and apricot are also recorded hosts, and (Glass and Fiori, 1955), orchardists have
damage can be significant in some situations. switched to other broad-spectrum insecti-
Differences in the host preference, develop- cides. Development of resistance to other
ment, diapause, phenology and population insecticides has occurred, although it has not
dynamics have been found for strains or always occurred in all countries or been doc-
races of the moth taken from apple, plum or umented adequately.
Organophosphates were the next chemi- attacking by predation (Knight et al., 1997) or
cal group used in many countries (azinphos- parasitism (Hassan, 1989) of eggs and
methyl, phosmet, diazinon and phosalone), neonate larvae (MacLellan, 1972). The cryp-
but resistance is now widely recorded tic habit of the larval stages (including dia-
(Barnes and Moffitt, 1963; Bush et al., 1993; pause) offers some protection against natural
Varela et al., 1993; Blomefield, 1994; Knight et enemies. In some situations, bird predation
al., 1994). Pyrethroids (bifenthrin, cyfluthrin, of diapausing larvae can be significant
cypermethrin, deltamethrin, esfenvalerate, (Wearing and McCarthy, 1992). However, the
fenpropathrin, fenvalerate, flucythrinate, flu- high levels of damage typically observed in
valinate and permethrin) have seen some the absence of controls indicate that biologi-
acceptance in the eastern USA (primarily for cal control, if present, is insufficient to main-
leaf-roller control), although the trend in tain the pest below the economic threshold,
much of Europe has been to avoid such which is relatively low.
broad-spectrum insecticides due to their neg-
ative impacts on natural enemies. MATING DISRUPTION The release of sufficient
In Europe, more selective insecticides synthetic sex pheromone to delay or prevent
have been increasingly used, including mating and provide control of codling moth
juvenoids (such as fenoxycarb (Charmillot, has been researched extensively worldwide,
1989)), chitin synthesis inhibitors (difluben- based on promising results with a range of
zuron, triflumuron, chlorfluazuron and formulations (Charmillot, 1978; Moffitt and
teflubenzuron) and ecdysone agonists (e.g. Westigard, 1984; Gut et al., 1992; Minks and
tebufenozide and methoxyfenozide) (Heller van Deventer, 1992; Judd et al., 1997). The
et al., 1992). However, there are also exam- mechanisms by which disruption acts are not
ples of resistance to these compounds entirely clear (Minks and Cardé, 1988) and it
(Moffitt et al., 1988; Sauphanor and Bouvier, may be possible to use pheromone-related
1995). In addition, avermectin (a macrocyclic compounds to improve results (Witzgall et
lactone fermentation product) has some effi- al., 1996a).
cacy (Cox et al., 1995), as does spinosad, Mating disruption is inversely density-
another fermentation product. The advan- dependent and therefore works best at low
tage of more selective insecticides is the pest densities in sites without significant
reduced impacts on natural enemies, per- immigration. It is not as effective in situa-
mitting the maximum contribution of bio- tions where the pheromone cloud is difficult
logical control against other pests. to maintain (steep slopes, windy sites, miss-
Petroleum oils have been used as ovicides ing trees or uneven orchard canopy) or in
(Webster and Carlson, 1942), although ear- close proximity to unmanaged populations.
lier products often caused phytotoxicity. The first commercially available pheromone
More recently, highly refined and purified dispenser for control of codling moth
products have been shown to have good (Isomate-C®) became available in the USA in
efficacy (Riedl et al., 1995) and have reduced 1991. Mating disruption of codling moth is
phytotoxicity problems. Particle films now commercially accepted in several coun-
(Unruh et al., 2000) also have some efficacy tries, and c. 40,000 ha of orchards were
against codling moth. treated with pheromone formulations in
Mechanical control, using bands on tree Washington, California and Oregon in 2000
trunks to collect diapausing larvae, has also (G. Thayer, Oregon, 2000, personal commu-
been used, but these do not collect the pro- nication). This has occurred in part because
portion of the population that falls to the of the failure of conventional insecticides,
ground directly. They can be effective if due to resistance, as well as the intrinsic
used in conjunction with other tactics (e.g. environmental and worker safety of
Judd et al., 1997). pheromone products.
Although codling-moth mating disrup-
BIOLOGICAL CONTROL There are a range of tion is not yet registered in all European
biological control agents of codling moth, countries, it has been widely used in some
areas (e.g. northern Italy). The relatively Europe (Audemard et al., 1992), including the
higher cost of this technique slows its adop- UK (Glen and Payne, 1984), New Zealand
tion, especially in warmer regions where two (Wearing, 1990) and the USA (Westigard and
applications per season of the dispenser are Hoyt, 1990). In hot climates with high levels
necessary. of solar radiation, the persistence of the virus
in the field is poor (about 1 week), making
MASS TRAPPING AND ATTRACTICIDAL CONTROL frequent applications necessary. However, its
Mass trapping has not proved to be very effectiveness against high pest populations,
effective against codling moth (e.g. Proverbs in combination with mating disruption,
et al., 1975), in part because of the cost and offers organic apple growers an effective way
practical difficulties of deploying sufficient of reducing pest populations to levels at
stations. As with mating disruption, the tac- which mating disruption can operate effec-
tic aims to prevent mating and therefore tively. Commercial use of the virus has unfor-
pest progeny. However, whereas in mating tunately been limited by the costs of
disruption males can survive to find a mate production using live insects. Industrial-scale
the next night, this is not possible where production offers reduced costs to growers
males have been removed from the system, (M. Guillon, personal communication), which
which represents a potential strength of the should assist adoption in future.
approach. If droplets containing sex
pheromone and a fast-acting insecticide are
19.3.2.2 Oriental fruit moth and other
used instead of traps (Charmillot et al.,
Grapholita (= Cydia) species
1996), then the costs can be somewhat
reduced. It may also be possible to develop Grapholita molesta (Busck) (Plate 19.4) and
multiple-species attracticides (Suckling and other members of the Grapholita genus, such
Brockerhoff, 1999). as G. lobarzewskii (= Cydia lobarzewskii) and G.
janthinana (Cydia janthinana (Dup.)) are some-
STERILE-INSECT TECHNIQUE Although it is tech- times recorded as pests of apple (Kalman et
nically feasible (e.g. Proverbs et al., 1982), al., 1994). In several countries, G. molesta (or
sterile-insect release is expensive and has oriental fruit moth) is reported to be increas-
several important limitations. Most impor- ingly important as a pest of apples (e.g. Reis
tantly, it requires mass rearing with special- et al., 1988; Pollini and Bariselli, 1993). These
ized capital-intensive facilities, excellent species typically feed on shoots early in the
quality control to maintain mating competi- season, as well as fruits later in the season.
tiveness with feral insects, geographical iso- The biology and options for control are simi-
lation, political support and ongoing lar to those for codling moth, but the pest
investment in the event of movement of con- status may not always warrant intervention.
taminated fruit. There are apparently few Within the past few years, oriental fruit moth
regional orchard industries that meet these has emerged as a major pest in several mid-
criteria. A sterile-insect release programme western and eastern US growing districts,
was commenced in the 1990s to eradicate the surpassing codling moth in importance.
codling moth from the 8000 ha of apple and
pear trees in the Okanagan valley in British
19.3.2.3 Tephritid fruit flies (Diptera:
Columbia. While successful in some respects,
Tephritidae)
the goal of eradication has not been realized
and the programme has been redirected to a True fruit flies of the family Tephritidae (Aluja
minimal-insecticide control programme (H. and Norrbom, 2000) deposit eggs directly into
Thistlewood, personal communication). the flesh of developing fruit, particularly fruit
approaching readiness for harvest. The tiny
MICROBIAL CONTROL The most promising puncture made through the skin of fruit dur-
microbial control against codling-moth ing egg-laying is difficult to detect without
neonate larvae is a granulosis virus (Tanada, magnification and may remain so even when
1964), which has been tested extensively in underlying flesh has decayed substantially
during larval feeding. Commonly, infested orchards, annual invasion by adults from
fruit are detected only after a few days of beyond orchard perimeters represents a
exposure to room temperature following pur- major challenge to managing these pests. In
chase by an unwary consumer. many situations, not owning the land beyond
Three species of tephritid flies are key orchard perimeters severely compromises
pests of apples in geographical areas where growers’ ability to reduce invading flies
their presence coincides with commercial through eliminating nearby unmanaged host
apple production. The apple maggot fly, R. trees. This may be especially problematic
pomonella (Walsh) (Plate 19.5), is native to where orchard blocks are comparatively
North America and is not known to occur small and perimeters are exposed to consid-
elsewhere. It is especially important as a pest erable non-orchard vegetation.
of apples in eastern and midwestern regions. Currently, all three pest species are man-
It has a more limited, but growing, distribu- aged primarily by applications of
tion in the western fruit-growing regions. organophosphate insecticides, although in
The Mediterranean fruit fly, Ceratitis capitata some areas the preharvest interval dictates the
(Wiedemann), is native to Africa and has use of pyrethroids. Applications are timed in
spread to most fruit-growing regions of the accordance with the occurrence and abun-
world. It has become an important pest of dance of captures of invading adults by moni-
apples in Middle Eastern countries, includ- toring traps placed on perimeter trees.
ing Israel, Syria and Turkey. The South Predictive phenology models (Jones et al.,
American fruit fly, Anastrepha fraterculus 1989) have been useful in determining the
(Wiedemann), is native to South America but timing of emergence. In some cases, confining
has spread to Central America and Mexico. insecticide application only to perimeter trees
Recently, it has begun to damage commer- or baiting perimeter trees with odour–visual
cially produced apples in southern Brazil traps has provided effective control (Cohen
(Sugayama et al., 1998). For all three species, and Yuval, 2000; Prokopy et al., 2000). Even
there is an extremely low tolerance, border- though there are no known cases of insecti-
ing on zero, for larval-infested fruit, espe- cide resistance in any tephritid fly, the need
cially fruit intended for export. for continuous protection of apples by insecti-
Sometime during the past two centuries, cide residue over the course of the 2–3-month
all three species expanded their host range to period of susceptibility to fly oviposition is
include apples. In the process, they have prompting some growers to seek alternative
escaped most of their natural enemies (partic- approaches to fly management.
ularly parasitoids), which provide some bio-
logical control of fruit-fly eggs or larvae in
19.3.2.4 Leaf-rollers (Lepidoptera: Tortricidae)
native host fruit. Apparently the chemical
and physical properties of apples are suffi- BIOLOGY Leaf-rollers have only an indirect
ciently similar to those of the native hosts of physiological impact on the tree, since they
these flies to have facilitated fly colonization feed on the fruit surface rather than the
of apples but are different enough from the seeds. While the impact on the tree may be
native hosts to exclude colonization by para- negligible, the impact of fruit feeding on
sitoids, most of which respond only to highly grower returns is a direct one. Leaf-rollers
specialized cues when searching for hosts. In emerge as a major concern in many orchards
consequence, fly populations on feral or oth- that apply selective controls for codling
erwise unmanaged apples or other newly moth, as well as for exporters forced to meet
acquired host trees can build to large num- quarantine tolerances with a nil threshold.
bers and severely threaten apple orchards Larvae typically web foliage together and
within a kilometre (in the case of apple mag- many also feed directly on the fruit surface.
got fly) or more (in the case of Mediterranean This cryptic habit has often made insecticidal
and South American fruit flies). Despite control difficult. Fruit damage is visible as
grower vigilance in preventing fly oviposi- scarring or corking or as rots associated with
tion and larval development in commercial open wounds in storage, and larvae occa-
sionally enter the apple calyx. Injury to fruits (florists’)), Crataegus (hawthorns), cotoneaster,
destined for fresh and especially export mar- Eucalyptus, Humulus lupulus (hop), Jasminum
kets has the most significant economic (jasmine), Ligustrum vulgare (privet), Litchi chi-
impact, compared with that of processing- nensis (lychee), Macadamia integrifolia
grade apples. (macadamia nut), Medicago sativa (lucerne =
Leaf-roller biology differs in several alfalfa), Pinus (pines), Prunus persica (peach),
important ways from the internal feeding Prunus armeniaca (apricot), Pyrus (pears),
tortricid species (van der Geest and Quercus (oaks), Rubus (blackberry, raspberry),
Evenhuis, 1991). Many have much wider Solanum tuberosum (potato), Trifolium (clovers),
host ranges and feed on leaves as well as Vicia faba (broad bean), Vitis vinifera
fruit (Chapman and Lienk, 1971). Their (grapevine), Ribes (currants), Rosa (roses), cit-
external life habit is accompanied by larval rus, Diospyros (malabar ebony), Populus
dispersal through ballooning, typically fol- (poplars), Vaccinium (blueberries).
lowed by the establishment of a larval nest
on shoots or the undersides of leaves. Larger DETECTION AND INSPECTION METHODS Phero-
larvae are able to relocate to fresh nests and mones are known for many tortricids
use their silken thread for both nest con- affecting apples (http://www.nysaes.
struction and escape. Many species are mul- cornell.edu/pheronet), and traps have been
tivoltine, with up to four generations per widely used for detection and monitoring of
year. Unlike codling moth, few leaf-roller leaf-roller populations. A range of applica-
species are geographically widespread. tions were reported by Suckling and Karg
Instead, apple-growing regions typically (2000), including species-distribution sur-
have a unique complex of leaf-roller species veys, insecticide-resistance monitoring,
(Table 19.2; Chambon, 1986). insecticide spray-reduction programmes and
sample collection for population studies.
HOST RANGE Many leaf-rollers attacking apple More labour-intensive systems involving lar-
have very wide host ranges. The following val assessments on shoot tips have also been
represents a partial list of hosts of the light used for predicting the size of subsequent
brown apple moth, Epiphyas posvittana, to indi- generations within a season.
cate the range of economically important alter- Modern diagnostic methods are also under
native hosts: Actinidia chinensis (kiwifruit), development for a range of tortricids. Several
Chrysanthemum morifolium (chrysanthemum DNA methods have been used for species
Table 19.2. Abbreviated list of leaf-roller pests affecting apple in various regions.
Species Common name Distribution
Adoxophyes orana (Fischer von Summer-fruit tortrix Europe, Asia

Röslerstamm)
Archips argyrospila (Walker) Fruit-tree leaf-roller North America
Archips breviplicanus (Walsingham) Asiatic leaf-roller Asia
Archips podana (Scopoli) Great brown-twist moth Europe, Asia
Archips rosana (L.) European leaf-roller Europe, USA
Archips xylosteanus (L.) Apple leaf-roller Eastern Europe
Argyrotaenia velutinana (Walker) Red-banded leaf-roller Eastern USA
Choristoneura rosaceana (Harris) Oblique-banded leaf-roller North America
Epiphyas postvittana (Walker) Light brown apple moth Australia, New Zealand
Pandemis heparana (Denis and Schiffermüller) Pandemis leaf-roller Europe
Pandemis limitata (Robinson) Pandemis leaf-roller North America
Pandemis pyrusana Kearfott Pandemis leaf-roller Western USA
Platynota flavedana Clemens Variegated leaf-roller Eastern USA
Platynota idaeusalis (Walker) Tufted apple-bud moth Eastern USA
identification (Sin et al., 1995; Gleeson et al., lineatus Barber; Hempitera: Rhopalidae) has
2000), and this approach should provide ready similar pest status. Damage usually occurs in
taxonomic support for ecological studies. the latter part of the season and is character-
ized by a spongy, depressed area c. 1 cm in
BIOLOGICAL CONTROL Reduction in broad- size surrounding the feeding puncture.
spectrum insecticide use on apple is typically Externally, damage can resemble physiologi-
accompanied by an increase in biological- cal disorders such as bitter pit, but the tissue
control activity of leaf-rollers and other pests. beneath the skin does not turn brown.
An example is the spread of the parasitoid
wasp Colpoclypeus florus Walker (Plate 19.6)
19.3.2.7 Miscellaneous opportunists
for control of the oblique-banded leaf-roller,
Choristoneura rosaceana (Harris) (Plate 19.7) A number of insects are attracted to ripening
after the reduction of organophosphate use in or overripe fruit and will either create a
Washington. In many cases, leaf-roller para- point of entry or enlarge damage due to
sitoids and predators are present on alterna- other causes (splits, stem punctures, etc.).
tive host plants outside orchards and follow Vespid wasps are often found in orchards
the pest populations across a range of host near harvest and, although they are primar-
plants (Suckling et al., 1998). A fuller treat- ily predacious, they chew holes in ripe fruit
ment of leaf-roller biological control is pre- and pose a hazard to harvesters. Nitidulid
sent in Mills and Carl (1991). beetles are also attracted to ripening fruit
and can be found feeding under the surface.
Earwigs are orchard residents that are usu-
19.3.2.5 Cutworms and fruit worms
ally predacious, but will also chew or
(Lepidoptera: Noctuidae)
enlarge holes in fruit. They can curl up in the
Although minor in importance in compari- stem cavity and make their way into the
son with the tortricids, several species are packing-house. The dock sawfly, Ametastegia
capable of fruit feeding later in the season. glabrata (Fallén), tunnels into the fruit, espe-
The larvae excavate shallow round holes in cially those close to the ground, in order to
the fruit, rendering them unmarketable. The find an overwintering shelter.
spotted cutworm (Xestia c-nigrum (L.)),
Bertha army worm (Mamestra configurata
Walker), variegated cutworm (Periodroma 19.4 Foliage Feeders
saucia (Hübner)), black cutworm (Agrotis
ipsilon Hufnagel) and the western yellow- There are multiple groups of arthropods that
striped army worm (Spodoptera praefica) are a attack and feed mainly on foliage, with the
few of the species that can damage apple primary damage being loss of photosynthetic
fruits and leaves. More recently, a new capacity due to loss of chlorophyll and dis-
species, Lacanobia subjuncta (Grote & rupted osmotic balance. From the perspective
Robinson), was recorded from Washington of plant productivity, specifically yield para-
State (Landolt, 1998) and has become an meters, the effect of chlorophyll loss is con-
important pest in some areas. troversial. No clear and uncontested
relationships have been established, although
it seems clear from the body of literature that
19.3.2.6 Fruit-stinging insects (Hemiptera)
there is not a directly proportional relation-
Pests in this group are also orchard invaders ship between loss of chlorophyll and loss of
and damage levels are often highest around photosynthetic capacity.
the orchard borders. The surrounding habi- Trees are capable of sustaining a certain
tat is a primary determinant of the intensity degree of foliar damage without any mea-
of attack. The most common example are the surable loss in yield; thus, the critical ques-
stinkbugs (Pentatomidae; Euschistus consper- tion becomes: ‘How much damage?’ The
sus Uhler and Acrosternum hilare (Say)), but studies performed attempting to establish
the western box-elder bug (Leptocoris rubro- such relationships quantitatively have been
restricted in interpretation to the particular nensis, Eotetranychus carpini, Bryobia spp.)

combination of cultivar, climate and growing cause a similar type of damage, but are
regime in which they were conducted and, regional in distribution. A few species of
as a consequence, the results and interpreta- tenuipalpids (false spider mites) and tarson-
tion have been quite variable. emids (e.g. Cenopalpus pulcher Canestrini &
The implication of some level of tolerable Fanzago) are apple pests in some regions
damage has a critical implication for IPM: (Jeppson et al., 1975b).
the latitude for biological control. In many The biological control of spider mites is
cases, some level of the pest population must well studied and implemented, with vary-
survive in order for the natural enemy to ing degrees of success. The predatory mites
survive (unless there is an alternative host). in the family Phytoseiidae (Kostianinen and
Unlike pests of quarantine importance or Hoy, 1996) are the most frequent and suc-
direct fruit feeders, there is a greater window cessful biological-control agents (e.g.
of opportunity for non-pesticidal control Typhlodromus (= Galandromus = Metaseiulus)
measures, since the need for control is not so occidentalis (Plate 19.10), Typhlodromus pyri
immediate or triggered at a low threshold. Scheuten, Amblyseius fallacis (Garman),
Given the societal emphasis on reduction in Ambylseius andersoni (Chant), Neoseiulus cali-
pesticide use, this characteristic should be fornicus (McGregor)) (Jeppson et al., 1975a).
more fully exploited in the future. Different species are better adapted to differ-
Many of the foliage feeders are classed as ent growing regions; for example, T. occiden-
secondary (in importance) or induced pests. talis is ideally suited to the arid climate of
The latter classification implies that they the western USA, whereas T. pyri requires a
would not have achieved pest status with- more humid, temperate climate (Beers et al.,
out pesticide inputs directed at a primary 1993). T. pyri is the most important mite
(often direct) pest. Again, this points to the predator in the temperate regions of
opportunity to regulate this group using Europe (excluding Scandinavia and the
non-pesticidal methods, or only on an occa- Mediterranean region). It is widely used for
sional basis. European red-mite control, often through
the release of organophosphate (OP)-
resistant strains (Blommers, 1994). The
19.4.1 Mesophyll stylet feeders number of years needed to achieve success-
ful spider-mite control may vary between 1
This group feeds on cellular contents (temperate conditions) and 3 (cooler con-
(including chlorophyll) by penetrating the ditions). T. pyri does not occur in the
leaf surface (often from the underside), Mediterranean area, where summers are too
killing only one or a group of cells at each hot and dry. A. andersoni is the most impor-
feeding site. The damage appears as speck- tant predator in these areas, where its nat-
ling (leafhoppers) or bronzing (tetranychid ural populations can very successfully
and eriophyid mites), depending on the size control the pest populations (García-Marí et
of the mouth-parts and the depth of penetra- al., 1989).
tion. Reduction in photosynthesis may fol- Several predatory mite species have
low extensive feeding, due possibly to a adapted well to orchard spray regimes, and
combination of chlorophyll loss and/or this is, in large part, the reason why inte-
stomatal closure caused by water loss. grated control programmes have been possi-
ble (Hoyt, 1969). In addition, several species
are reared commercially and sold for release
19.4.1.1 Tetranychid (spider) mites
in orchards either as an inoculative measure
Several species are worldwide pests of or as a sort of ‘living pesticide’; some strains
apple, including the European red mite (P. have been selected for tolerance to pesticides
ulmi (Koch) (Plate 19.8) and two-spotted (Hoy and Knop, 1981; Roush and Hoy, 1981).
spider mite (T. urticae Koch) (Plate 19.9). Other families also contain predatory species
Other species (T. mcdanieli, Tetranychus vien- useful in apple orchards (e.g. Trombidiidae,
Anystidae, Stigmaeidae and Tydeidae); how- with no exposure on the leaf surface. The
ever, these predators usually play a support- entire preimaginal period is spent in the
ing role to the phytoseiids. In the mine, which is formed between the upper
mid-Atlantic area of the USA, a predatory and lower epidermis by the larva’s feeding
coccinellid (Stethorus punctum (LeConte)) activities. The shape of the mine is usually
provides the greatest degree of biological characteristic of the species or group: the
control, whereas a related species in the gracillariid leaf-miners (Phyllonorycter (=
western USA (Stethorus picipes Casey) plays Lithocoletis) blancardella, Phyllonorycter
only a minor role. Several groups of preda- elmaella, Phyllonorycter crategella), or so-
tory bugs (especially mirids in the genera called ‘tentiform’ leaf-miners, produce a dis-
Campylomma, Campyloneura, Blepharidopterus, tinctive dome-shaped mine with white
Atractotomus) will prey on mites and may feeding specks visible from the upper leaf
play an important role in biological control. surface. Two species of lyonetid moths
The relative dominance or contribution of a (Leucoptera malifoliella (= scitella) and Lyonetia
predator is governed by many factors, clerkella) produce a blotch and sinuous mine,
including climate, pest complex, surround- respectively (Alford, 1984). Several species
ing habitat and spray regimes prevalent in of coleophorid moth (case-bearers) also form
the area. mines, but these are usually minor pests.
The cryptic habit of the larvae presents
some challenges for chemical control. Either
19.4.1.2 Eriophyid mites
the adult must be targeted with applications
There are two basic groups of eriophyids, sufficient to cover the entire flight period or
free-living and those causing plant deformi- the pesticide must penetrate the leaf surface
ties (galls or blisters). In the former category, in order to deliver the toxicant to where the
the apple-rust mite, Aculus schlechtendali larvae are feeding. True systemic insecti-
(Nalepa), is widely distributed and com- cides are now rare and, because of residue
mon, but rarely considered a serious pest. problems, few are being developed.
While high populations can cause leaf Insecticides with translaminar activity are
bronzing and premature terminal bud set sufficient and typically present few prob-
(Hull et al., 1986), it is considered a quasi- lems in the registration process. While sev-
beneficial species in some areas in that it eral effective insecticides are registered for
provides an alternative prey for predatory use against leaf-miners, biological control
mites (Hoyt, 1969). Sensitive cultivars (e.g. has been reasonably well studied and par-
‘Golden Delicious’) may be russeted by tially implemented. Parasitic wasps (e.g.
feeding in the calyx area, which occurs Pnigalio flavipes, Pnigalio marylandensis,
shortly after bloom. Examples of the gall- Apanteles ornigis) are regionally abundant
formers attacking apple are few. Burts (1970) and can provide substantial levels of con-
reported on two closely related species trol. However, hymenopterous parasitoids,
Eriophyes (Phytoptus) pyri (Pagenstecher) and as a group, tend to be less tolerant of broad-
Eriophyes mali (Burts), both of which may spectrum insecticides, and biological con-
attack apple. They cause blisters on the trol is easily disrupted.
leaves and fruit, leaving the latter scarred
and deformed. The current spray pro-
19.4.1.4 Skeletonizers
gramme has made these mites rare.
There are several species of arthropods from
various groups that skeletonize leaves, but
19.4.1.3 Leaf-miners
none are specialists on apple and their signif-
Several families of microlepidoptera (moths) icance is sporadic and local. Examples
mine apple leaves in the larval stage. The include the apple and thorn skeletonizer
egg is laid on the surface of the leaf (usually (Eurtomula pariana; Lepidoptera: Choreutidae)
the underside) and the newly hatched larva and the pear slug (Caliroa cerasi; Hymenoptera:
penetrates the leaf directly from the egg, Tenthridinidae).
19.4.2 Bulk leaf feeders (Quadraspidiotus perniciosus (Comstock)) is

widely distributed and, left unchecked, can
This is a varied group, comprised mostly of cause reduced tree vigour or even mortality
polyphagous Lepidoptera. Many are pests of (Plate 19.11). Scales feed primarily through
deciduous forest trees, which can use apple tree bark, forming large encrustations that
as a host in the absence of pesticide residues. devitalize the tree. Mealybugs (especially
Examples include the notodontid moths Pseudococcidae) also suck plant juices, but
Datana ministra and Schizura concinna and usually choose more tender tissues (shoots
several species in the lasiocampid/ and leaf axils) as feeding sites. In the latter
lymantriid group (Orygia antiqua; Euproctis case, the primary damage is not from
chrysoheoea, brown-tail moth; Euproctis removal of plant product, but rather the pro-
similis). The winter moth (Operophthera bru- duction of honeydew (liquid drops of excre-
mata) is occasionally an important pest of ment rich in simple sugars). Honeydew
apple in Europe. The autumn webworm dripping on fruit can cause fruit russeting on
(Hyphantria cunea; Arctiidae) is an example of sensitive cultivars or can support the growth
a gregarious nest maker, which forms a large of sooty mould, a superficial but unsightly
web (up to 50 cm long) and devours all leaf fungal growth.
material inside it. Other gregarious lepi- Both scales and mealybugs are considered
dopterans include the tent caterpillars to be induced secondary pests, which would
(Malacosoma americana, Malacosoma fragilis occur only at low levels if their natural-
and Malacosoma disstria; Lasiocampidae) and enemy complex were not decimated by
the ermine moths (Yponomeutidae, e.g. broad-spectrum pesticides. Currently, the
Yponomeuta malinellus (apple ermine moth)); pre-bloom use of horticultural spray oils
and others in the genera Swammerdamia and appears to keep scales in check, although this
Paraswammerdamia are capable of using apple activity is probably supplemented by in-sea-
as a host. Currently, these are primarily pests son use of organophosphates. Mealybugs, on
on unsprayed back-garden trees, but they the other hand, can be extremely persistent
represent a rich pool of potential insect once established (usually in large, older trees)
species that may respond to our changing and even an intense spray programme can
pest-control regimes. only keep them in check, not eradicate them.
Both species will infest the fruit towards the
latter part of the season, especially when
19.5 Structural Feeders populations are high. A red ring appears
around the scale that settles on fruit; mealy-
The group is defined as those attacking plant bugs usually move to the calyx, where detec-
parts other than fruits and foliage, that is, tion is difficult during packing operations.
branches, trunk and root systems. The group Feeding in the calyx end causes a softening
is a varied one taxonomically, and several of and deterioration, which may be exacerbated
the pests included cross the damage-classifi- by long-term storage. Quarantine measures
cation boundaries as defined here. While and food contamination are issues with these
some of these pests can cause sufficient dam- two groups of pests.
age to cause tree death, as a group they are The psyllids (Homoptera: Psyllidae) are
generally considered less important than the key pests of pear, but one species, Psylla mali
fruit and foliage feeders. (apple sucker), is a corresponding pest of
apple in some regions. Like pear psylla, this
pest feeds on shoots and produces honey-
19.5.1 Superficial woody-tissue and shoot dew, with the attendant problems for fruit
feeders and vegetative growth. However, its impor-
tance on apples is minor in magnitude com-
Two groups of Homoptera (scales and pared with the related species attacking pear.
mealybugs) are widespread and sometimes Another large and important group of
important pests of apple. San Jose scale homopterans (aphids) may also be classed as
shoot feeders. This group has specialized in insertum) are occasional pests of apple,
phloem feeding and is a large and successful using one of the Gramineae as their summer
group of pests on many crops. The aphids host. Only very heavy infestations, which
that feed on apple may use it as their only can infest developing fruitlets, are consid-
host or as the primary or overwintering host, ered damaging.
with a different plant species as a summer Aphids have a rich and varied natural-
host. The two life-history patterns have a enemy complex that prey on them, including
definite influence on management. lacewings (Chrysopa and Hemerobius), coc-
Apple aphid (Aphis pomi) (Plate 19.12) is cinellids (ladybirds), various parasitic wasps
a widespread pest of apple, occurring in and a variety of predatory mirids (e.g.
most apple-growing regions of the world. It Campylomma, Deraeocoris, Orius). Despite
spends its entire life cycle on apple, repro- this, aphids often escape from biological con-
ducing by parthenogenesis for the greater trol. Many of their predators are generalists
part of the season. Winged (alate) forms are and will only be attracted to large aphid
produced under high population levels to populations (i.e. after the point where con-
colonize new host plants, and in the trol is needed or desired). A number of
autumn sexual forms are produced, which broad-spectrum pesticides used in orchards
mate and lay overwintering eggs. Both are toxic to one or more of these natural ene-
leaves and shoots are attacked, and some mies and disruption of biological control
level of growth reduction is assumed under early in the season may preclude stable regu-
heavy attack; however, most of the concern lation for the rest of the season.
for this pest involves the production of hon- Woolly apple aphid (Plate 19.13) was one
eydew and sooty mould and the resulting of the earliest targets (1920s and 1930s) of a
fruit contamination. widespread introduction of a biological-con-
Several other common aphid species are trol agent, the parasitic wasp Aphelinus mali.
heteroecious, although their damage may This wasp was introduced in many of the
be quite distinct from that of apple aphid. areas around the world where woolly apple
The rosy apple aphid (Dysaphis plantaginea aphid had also been introduced and was
or Dysaphis devecta) also feeds on shoots and successfully established in most areas
leaves, but injects a salivary toxin, which (Yothers, 1953). It is still thought to provide
severely deforms both organs. In addition, the primary means of biological control
the toxin causes fruit deformity on sensitive today, although the generalist predators
cultivars. This pest colonizes a herbaceous described above also play a role.
weed host during the summer; thus control
measures must occur fairly early in order to
be effective. Woolly apple aphid uses elm as 19.5.2 Wood-boring insects
the alternative host in some areas, but is
functionally monophagous in the north- Several families of Lepidoptera attack the
western USA and Europe. This species pro- cambium of the trunk and major scaffold
duces both aerial and edaphic colonies; the limbs, and prolonged attack can girdle and
former are easily controlled, the latter with kill these organs. The clearwing moths
great difficulty. The root colonies are (Sesiidae) have several species that attack
thought to devitalize the tree and, even various fruit and ornamental trees and at
though rootstocks were developed specifi- least one species that infests apple
cally to be resistant to woolly apple aphid (Synathedon myopaeformis; UK and continen-
(the Malling–Merton series), there is evi- tal Europe). One species of tortricid moth
dence that this resistance is breaking down. (cherry-bark tortrix, Enarmonia formosana)
Feeding by both the root and shoot colonies causes a similar type of damage.
produces galls; typically, the above-ground While rarely a problem in sprayed
galls (which occur in leaf axils) are pruned orchards, these insects can be difficult to con-
off and are of little significance. Several trol once established. It is difficult, if not
species of Rhopalosipum (R. fitchii and R. impossible, to kill larvae in their galleries
with pesticides; thus pesticidal control mea- because of groundwater contamination

sures must be directed at the adults. Typically issues. While biological-control agents are
they are univoltine, with a prolonged flight, known, their management is little studied
making continuous coverage a necessity. or applied. Entomophagous nematodes
The scolytid beetles comprise some of the (injected into the soil) may alleviate the prob-
more serious forest pests, and several species lem, but their effect is not well studied in
in the genera Scolytus and Xyloborus are pests tree fruits.
of apple. The larvae form distinctive galleries One very large species of cerambycid bee-
in the wood, and adults often bore into tle (Prionis sp.) can attack apple; control mea-
shoots just below buds, causing weakening sures are similarly difficult. Weevil
and breakage. In general, these insects are (curculionid) larvae are known to attack the
usually attracted to trees that are already root system on occasion, but the extent of
weakened by some other pest or disease, damage is not well defined. The adults of
although young trees can suffer damage some species may also be problematic when
when they are close to a source of emerging they feed on fruits, fruit stems or foliage.
adults. Coleopteran wood-borers in the fami- Woolly apple aphid is the only truly ubi-
lies Buprestidae and Cerambycidae may also quitous root pest of apple (see above),
attack apple, but are rare in sprayed com- although typically only the aerial colonies
mercial orchards. are treated.
19.5.3 Root-system pests 19.6 Conclusion
This is a fairly restricted group of pests, Apple pest management is continually

which are given little attention either evolving in response to changing horticul-
because they cause only occasional damage tural practices, the genetic structure of insect
or because of their cryptic life history. The populations, the importation or re-emer-
larvae of scarab beetles (several species in gence of new pests and societal pressures.
the genera Polyphylla and Pleocoma) feed on These pressures encompass fewer and safer
roots and can be locally severe. Trees on residues on food products, reduced environ-
sandy soils where the grubs thrive may suf- mental impact and the concept of sustainable
fer long-term damage; orchardists will often agricultural production. The result has been
replant repeatedly, trying various combina- increased regulation of pesticide use world-
tions of fertilizer or watering to promote tree wide and incentive programmes (specialty
growth, when in fact the root system is being labels) that promote reduced-impact pest-
systematically destroyed. management programmes. With the global-
Soil fumigation is currently the best rem- ization of fruit marketing, it is likely that all
edy to allow trees sufficient time to establish countries wanting to export apples will have
before the beetles reinfest the orchard. Soil- to conform to production and pest-manage-
applied pesticides are widely discouraged ment practices that embrace these concepts.
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20 Apple-orchard Freeze Protection
Schuyler D. Seeley and J. LaMar Anderson

Plants, Soils, and Biometeorology Department, Utah State University, Logan, Utah,
USA
20.1 Site Selection 522

20.2 Cultural Practices 523
20.2.1 Soil 523
20.2.2 Orchard canopy 523
20.2.3 Tree condition 523
20.3 Energetics of the Orchard Environment 524
20.3.1 Radiation 524
20.3.2 Conduction 524
20.3.3 Convection 524
20.3.4 Latent heat 525
20.4 The Daily Thermal Cycle 526
20.5 Energy Changes in the Orchard Canopy 528
20.6 Water in the Orchard Atmosphere During Freezes 528
20.6.1 Relative humidity 529
20.6.2 Dew-point/frost point 529
20.7 Freeze Types 531
20.8 Monitoring Freezes 531
20.9 Methods for Freeze Protection 533
20.9.1 Heating 533
20.9.2 Overhead irrigation 534
20.9.3 Under-tree sprinkling 535
20.9.4 Wind machines 535
20.9.5 Bloom delay 536
20.10 Summary 538
Apple production on a per-hectare basis has such a severely competitive business envi-
increased greatly over the past few decades. ronment, the successful apple grower must
Earlier-producing, higher-yielding, high- produce the maximum yields of quality fruit
density orchards of newer cultivars have possible per hectare at the lowest possible
made apple production a highly competitive cost in order to survive. In many fruit-pro-
business. Factors that have increased producing areas, losses from freezes are sub-
duction have resulted in decreased or nega- stantial. In fact, freeze damage losses in
tive profit margins due to excess supply. In apple orchards have exceeded the combined
522 S.D. Seeley and J.L. Anderson
losses to all pests. In this chapter we shall apples can be made from the accumulated
consider details of freeze protection for the temperature data. Freeze potential can then
apple orchard. be determined from the temperatures on
each site during bud break, anthesis and
full-bloom periods. Probabilities of advec-
20.1 Site Selection tive and radiative freezes can be developed
from a few years of accurate temperature
The best way to avoid freeze problems is to data that appropriately represent the char-
select a freeze-free site. Orchard-site visits acteristics of a particular site. Approx-
with veteran orchardists, area fruit agents imations of the climate of potential fruit
and professional pomologists can be very areas and long-term normals for existing
valuable for assessing freeze susceptibility. weather stations may be obtained from state
If apple trees are present, spur system climatologists.
anatomy can reveal their recent bearing his- Good sites may have problem subsites.
tory. Cluster-base enlargements that indi- Small valleys with no outlet for air drainage,
cate fruit-production history can be low spots in otherwise good orchard land
observed for the last 5–10 years. Spur sys- and obstacles to air movement, such as
tems should have such cluster-base enlarge- wind-breaks, swales, borders, basins and
ments every other year if the area is a good roadways, often cause cold-air stagnation
freeze-free site (Fig. 20.1). If a new fruit area and freezes. These freeze-liable subsites can
is to be developed, temperature measure- be used for alternative crops that tolerate
ment over a period of time is usually neces- low temperature during part of their produc-
sary to select a satisfactory site with tion cycle or used for roadways, drainage or
confidence. Inexpensive data-loggers can be equipment- and bin-storage areas.
placed in temperature-monitoring locations Other site considerations should be
in appropriate shelters on potential sites included in the analysis of each potential
and data can be downloaded monthly. fruit-orchard area. The soil should be rela-
Temperatures should be accumulated for at tively rock-free, of medium texture, of neu-
least 24 months – longer if possible – so that tral pH, of high water-holding capacity and
accurate estimations of chill, anthesis and fertile. The site is more important than the
growing degree-day accumulations can be soil because the soil may be modified to
determined. Estimation of bloom dates for advantage on a good site, but the best soil on
Fig. 20.1. Apple spur systems showing cluster-base enlargements where fruit were borne on every other
year’s growth in a productive orchard.
Apple-orchard Freeze Protection 523
a poor site will not allow profitable fruit pro- orientated to allow air drainage down-slope.
duction. Prevailing winds, often common in Channels should be maintained in natural
valleys and canyons, may help reduce freeze low areas to encourage cold air flow through
probability due to their ability to mix inver- such areas and away from the orchard. Small
sions. While windy sites may be advanta- trees also have the inherent problem of occu-
geous for freeze prevention, they may be a pying the coldest part of an inversion-tem-
hindrance to honey-bee activity, pollination, perature profile. Older orchards with taller
spray application and tree conformation. For trees and wider alleyways allow more air
additional information on the characteristics flow out of the orchard and occupy space
of a good site, see Chapter 11. higher in an inversion.
20.2 Cultural Practices 20.2.3 Tree condition
20.2.1 Soil 20.2.3.1 Irrigation
Fine-textured soils that are compact, full of Irrigation should be used to avoid tree stress
water and devoid of surface insulation have during the late summer and autumn when
greater heat adsorption and storage capacity tree maturation and acclimatization occur.
than medium- or coarse-textured soils, soils Inadequate irrigation will stress trees
with less water due to their lower water- through drought and excessive irrigation
holding capacity and soils with insulating will limit root respiration due to lack of oxy-
mulch layers on their surfaces. When gen. Irrigations should be timed to fill the
medium- or coarse-textured soils contain lit- root profile with adequate water for the
tle water and large amounts of air, they con- trees’ needs, but to avoid waterlogged soils
tain less heat and are poor conductors of the that limit root respiration and growth. Trees
heat they do contain. Often during winter, should not be stressed by limited water late
reoccurring freeze/thaw cycles cause soils to in the season to ‘harden’ them. Such stress
expand and develop numerous air spaces. will generally result in more limited cold-
Air spaces serve as insulators and decrease hardiness levels developing during the ensu-
conductivity. Firming the soil with irrigation ing winter season. After trees have matured
or other methods will decrease its insulation vegetatively and leaves have abscised, the
capacity. Other insulators at the soil/air soil should be filled to field capacity to
interface are weeds, mulches, grass cover ensure water availability through the winter.
crops, chopped prunings or rubbish. In drought conditions, winter irrigation is
Mowing or flailing machines can decrease advisable.
the effect of these insulative layers on the soil
surface and increase energy conductivity and 20.2.3.2 Nutrition
radiation. Uninsulated, water-filled, southerly-
exposed soils (in the northern hemisphere) Nutrition of apple trees should be main-
will accumulate significantly more energy tained at optimum levels during the growing
than insulated, air-filled, north-aspect soils. season. Nitrogen should be applied in early
spring, just before root growth commences,
and adequate supplies should be applied to
20.2.2 Orchard canopy give optimum shoot and fruit growth with-
out supplying excess. Avoid late applications
Modern high-density orchards of small trees that will delay late-summer maturation and
often incur problems with air flow. Trees that hardening of the trees. Growers are encour-
are planted close together impede the flow of aged to use foliar analysis of critical orchard
cold air through and out of the orchard. nutrients so that their fertilizer applications
Whenever possible, trees should be planted can be made in a timely manner and ade-
with the hedgerows or multiple-tree beds quately (see also Chapter 12).
20.2.3.3 Pruning and training This higher average temperature is mainly

due to the atmospheric absorption of
Pruning and training of apple trees should
energy. Other important selective absorbers
be done in the dormant season and in the
on the earth’s surface that contribute to
summer so that tree response will not pro-
temperature modification include ice, snow,
duce late-season, succulent, tender wood.
water vapour and carbon dioxide. A simpli-
The general rule is to avoid pruning from fied earth/atmosphere/solar energy bal-
dormancy induction in late summer through ance is given in Fig. 20.2.
the ensuing autumn and early winter period
(Plates 20.1 and 20.2).
20.3.2 Conduction
20.3 Energetics of the Orchard Conduction is energy transfer from mole-

Environment cule to molecule. A nail held in the hand
seems to be cool to the touch because iron
Energy is transferred in the orchard envi- has high heat conductivity. When a nail is
ronment by radiation, conduction, convec- held by the fingers on one end and in a
tion and latent-heat transfer. These flame on the other end, it soon becomes
processes are operative in any freeze event warm to the touch by molecular motion or
in the orchard and should be thoroughly conduction. Wood, as in a wood match, does
understood. not have a high heat of conductivity and can
support a flame on one end and still be held
by the fingers. Heat conductivity, or the abil-
20.3.1 Radiation ity to conduct heat by molecular motion, of
some common substances is presented in
Radiation is energy transfer by radiant Table 20.1.
energy or electromagnetic waves. Most
radiation in our solar system originates in
the sun. Radiation waves travel through 20.3.3 Convection
space and do not need molecules to facili-
tate their movement. This energy includes Convection is heat transfer by mass move-
infrared, visible and ultraviolet rays rang- ment of a fluid, such as oil, water or air.
ing from 1 millionth to 10 millionths of a Fluid particles having different tempera-
metre in length – often called short-wave tures and therefore different densities move
radiation. Short waves have greater energy freely to develop currents. On a sunny,
than longer waves, but they all travel at the warm, summer day, the major impact of the
speed of light. The earth also emits radia- sun’s radiation is received by the earth’s
tion, but, since the earth’s temperature of surface. The surface temperature rises dur-
288K (15°C, 59°F) is much cooler than the ing the day and air next to the surface
sun’s temperature of 6000K (5815°C, absorbs energy and expands by conduction
10,500°F), it emits weaker radiation 10 mil- of molecule-to-molecule energy from the
lionths to 20 millionths of a metre in length surface. This warmed, expanded air
– often called long-wave radiation. becomes more buoyant than the air above it
Through a balance between interception and rises. Cooler air displaces the warmed
and emission, the earth reaches a radiative air and in turn absorbs thermal energy and
equilibrium temperature. The earth is also rises. The cycle is repeated to set up
150,000,000 km (93,000,000 miles) from the convective currents of rising warm air,
sun and turns once a day, so its equilibrium called thermals. When horizontal motion
temperature without an atmosphere would occurs, the air movement is called a breeze
be about 255K (18°C, 0°F). However, with or wind. The appropriate term for horizon-
its atmosphere, the actual average earth- tally moving air with associated water
surface temperature is 288K (15°C, 59°F). vapour and other inclusions is advection.
The earth/atmosphere energy balance

Approximations - relative size of numbers indicates energy flux in and out of earth and atmosphere
Radiation out from atmosphere 64 6 Radiation out from earth surface

Clouds Clouds
Sunshine
24 24 20
Energy gain
(clouds)
6 110
Energy gain
Energy loss (atmosphere) (clouds)
Q
160 Energy gain (atmosphere)
Sunshine
Land and vegetation

Land & Vegetation
24 72 50 6 24 6 110
Land and vegetation

Water Water
Energy gain to earth due to: Energy loss from earth due to:
Radiation: Radiation to space
solar (direct) Evaporation
atmosphere (indirect) Conduction/convection to
Condensation atmostphere
Radiation to clouds
Energy gain by earth 146 Energy loss from earth 146
Fig. 20.2. Simplified earth/atmosphere/solar-energy balance. Numbers indicate magnitude of energy fluxes
in the system. Most important energy fluxes are indicated.
Table 20.1. Heat conductivity of selected

substances.
Conductivity
Material (cal s1 cm2 °C1)
Air at 20°C 0.00006

Dry soil 0.0006
Water and snow 0.0014
Wet soil 0.005
Ice (0°C) 0.005
Rocks 0.006
Iron 0.160
Silver 1.1
20.3.4 Latent heat goes into the water and becomes latent
heat. When the reverse happens, the latent
Latent heat is energy that is transferred heat is released into the environment. The
when there is a change of state. Water exists heat change for ice formation is called the
in solid (ice), liquid (water) and gaseous heat of fusion and amounts to 80 cal g1 of
(water vapour) states. When heat is applied water frozen. Thus, 80 calories are given off
to ice, the ice melts and the energy applied when the molecules in 1 g of water freeze
to become arranged with less molecular radiation – insolation that enters the earth
motion into ice crystals. As long as the and its atmosphere – only about 50%
ice/water phase transition occurs, heat is reaches the earth’s surface, some directly
evolved and the temperature of the system and some diffusely or scattered from the
will remain at 0°C. When all water is atmosphere. About 4–6% is reflected from
frozen, heat production ceases. The molec- the earth’s surface and 6% is reflected from
ular motion returns on thawing, with the atmosphere. When clouds are present,
energy absorption. A similar absorption or up to 20% is reflected by water vapour.
release of thermal energy occurs when Depending on atmospheric conditions, the
water evaporates (absorption) or condenses most important being water in clouds, up to
(release). The energy involved in the latent 20% is absorbed by the atmosphere.
heat of evaporation is the absorption of As the sun rises above the morning hori-
about 600 cal g1 of water. Conversely, zon, its energy is spread over a large area
when a 1 g of water condenses in the since it hits the round earth surface at a very
atmosphere, the latent heat of condensation low angle. As the earth turns, the impact
results in the release of 600 calories. The angle changes and the energy concentration
heat of condensation is 7.5 times the heat of reaching the surface per unit area increases
fusion! Large amounts of water are con- until solar noon. After noon, the reverse sce-
tained in the atmosphere, and the latent nario occurs until the sun sets below the
heat capacity of atmospheric water is an horizon. Thus, the energy curve through the
important source and sink for energy. The day resembles a cone (Fig. 20.3a). However,
evaporation of water from equatorial the temperature during the day depends on
regions removes energy from those areas. more than insolation. The earth, too, is radi-
Water-laden air moves advectively towards ating energy from its surface proportional to
the poles, where the water condenses and the surface temperature. Surface tempera-
releases energy. These processes are ture is highest in the early afternoon
tremendously important in the energy bal- because, even though the intensity of incom-
ance, advection and the prevailing winds of ing solar radiation decreases after noon,
the turning earth. They are also important incoming energy still exceeds outgoing sur-
in the orchard environment when a freeze face heat energy for a few hours. This
event occurs. energy contributes to the typical lag
between the time of maximum solar radia-
tion at noon and the maximum air tempera-
20.4 The Daily Thermal Cycle ture in an air thermometer several feet
above the ground in the afternoon (Fig.
The daily temperature cycle in the orchard 20.3b). At the time of the highest daily tem-
typically, but roughly, traces the thermal perature, the air temperature is highest at
influence of the short-wave insolation from the surface and decreases with increasing
the sun to the earth during the day and the elevation (Fig. 20.3c). After sunset, diffuse
long-wave radiation from the earth’s surface radiation energy has a small impact for a
during the night. During the day, an short time, but its effect is small when com-
insignificant amount of the total energy pared with the long-wave radiation from the
from the sun showers the earth and its earth’s surface.
atmosphere. But the magnitude is insignifi- Other insolation/radiative effects may be
cant only when compared with the total significant in the orchard microclimate.
energy emitted from the sun. This miniscule Hillsides, ledges, very large structures and
amount of the sun’s energy is highly signifi- large bodies of water may serve as reservoirs
cant to life on earth and is the source of for thermal energy and reradiate that energy
nearly all of the earth’s energy. In fact, the after sunset and during the ensuing night.
energy reaching the earth is sufficient to pro- Conduction of energy from the soil surface
duce the equivalent of 5000 t of coal s1 in or plant canopy to the adjacent air during the
photosynthate. Of the total incoming solar daytime warms the air and increases its buoy-
(a)
Noon Sunset Midnight Dawn
Relative air temperature
(b)
Convection Low Conduction
Energy exchange
energy to soil from air

radiation to air from soil
(c) Altitude (m) Relative temperature

200
In
D
ve
ec
Radiation inversion
rs
re
temperature curves
io
as wit
n cu
Ai
in h a
ai rv
rt
N
rt e
em wi
or
ai ltitu
em
m
rt d
pe th a
pe
al
em e
ra ltit
ra
ai
tu ud
pe
rt
tu
re e
ra
em
re
in
tu
Effect of wind machines

pe
cr
re
e
ra
as
tu
e
re
cu
rv
e
n
in versio
Top of
0
Add heat Effect of heaters
Noon Sunset Midnight Dawn

(d)
Warmer air
Co
ld
air
flo
w
Fig. 20.3. Energy changes in the orchard environment from solar noon until the next dawn. (a) Air-
temperature changes. (b) Incoming and outgoing radiation exchange, along with convection and
conduction of energy into the soil during the day and conduction of energy from the soil to the adjacent air
and outgoing radiation during the night. (c) Air-temperature profiles through the afternoon and night from
the surface to an elevation above the top of the developing inversion. Triangles show the effect of wind
machines and heating on the inversion-temperature profile. (d) Inversion created by cold air flow.
ancy. The warmed air rises in convective cur- thermal energy in buoyant air (due to less
rents through the daytime hours and into the dense air and lessening pressures at higher
night from thermal reservoirs. However, rising elevations) does not significantly affect the
normal temperature gradient from warmest at with cooler surface air. Inversions also form
the earth’s surface to lower temperatures at through the flow of cooler air near the
higher and higher elevations above the sur- ground into low areas (Fig. 20.3d).
face. Clouds of water vapour reflect solar radi-
ation into space and also reflect radiation from
the earth’s surface back towards the surface. 20.5 Energy Changes in the Orchard
The water-vapour content of the air is very Canopy
important in temperature changes, since it is
the main reservoir of atmospheric thermal Air does not hold as much energy as the
energy. During the day, all methods of heat water in the air. The molecular weight of
transfer are operating to develop the normal dry air (78% N2 , 21% O2 , 1% Ar plus small
daytime temperature gradient, with the high- amounts of other gases) is about 29 g (ª an
est temperature at the surface and decreasing ounce) and has a volume of 22.4 l (5.1
temperatures at increasing elevations. American gallons) at standard pressure and
Through the night, the surface of the earth temperature. Energy changes due to molec-
cools by radiation to space (Fig. 20.3c). In the ular motion of atmospheric gases are very
absence of short-wave incoming radiation and small on a molar basis, but, since the atmos-
as the surface cools, energy from the adjacent phere of the earth weighs about 5600
air and soil is transferred to the surface and trillion t (the weight of air above 1 ha at sea
this energy is, in turn, radiated to space. This level is approximately 104,000 t – equivalent
process continues during the night. The air to about 42,000 t acre1), energy changes
temperature near the earth decreases and over large areas and in large masses, such
reaches a low just after sunrise, when outgoing as the area over an orchard, become very
radiation still exceeds incoming radiation. It is significant. When water vapour is added to
important to note that the daily low tempera- the air and displaces the ‘average’ air mole-
ture occurs just after sunrise for this reason. cule, the volume of air containing water
The normal temperature gradient during becomes lighter because the molecular
the day is warmest at the surface and cooler weight of water (18 g) is 11 g mol1 lighter
at increased elevations above the earth’s than the average molecular weight of air
surface. During the night, incoming radia- (29 g). Therefore, water-laden air tends to
tion is very low and outgoing long-wave rise. More importantly, the energy-holding
radiation from the surface and conduction capacity of the more humid air is signifi-
of energy from relatively warm air near the cantly increased. At 25°C under normal
surface to the surface, with subsequent radi- pressure, water-vapour-saturated air holds
ation of that energy to space, set up a tem- 23 g m3 of water. A ‘box’ of such air above
perature gradient in the lowest few an orchard 100 m high would contain about
hundred feet of the atmosphere. The tem- 21,000 kg of water ha1 (~16,000 lb. acre1).
perature becomes cooler at the surface and Due to its energy content, especially during
increases with elevation to a higher temper- phase changes, water in the air is very
ature. Above the elevation with the highest important in energy exchanges between the
temperature, the temperature decreases environment and the plant. Energy relation-
again, as it did during the day. Since this ships of water in air are given in Table 20.2.
gradient near the surface is the inverse of
what it was during the day, the phenome-
non is called an ‘inversion’. An inversion 20.6 Water in the Orchard Atmosphere
results in a reservoir of warmer atmosphere During Freezes
above the orchard that may be located from
just a few feet to several hundreds of feet Water in the air stores large amounts of ther-
above the surface. The elevation of the mal energy compared with the air itself. The
warmest air is called the ‘top’ of the inver- water status of the atmosphere is described
sion. This reservoir of warm air can be used in several ways. The most important is rela-
in orchard freeze protection by mixing it tive humidity.
Table 20.2. Water status and energy exchange (arrows indicate changes in temperature or state of water).
Energy given off by water Energy taken up by water

Water vapour in hot air
+ 1 cal g1 °C1 Cooling ---------- Ø ≠ ----------- Heating 1 cal g1 °C1
Water vapour in cool air
Ê + 600 cal g1 °C1 Condensation ---Ø ≠------ Evaporation600 cal g1 °C1 ˆ
Ì
Deposition + 680 cal g1 °C1 680 cal g1 °C1 Sublimation
Ì
Water at the dew/frost point
Ë + 80 cal g1 °C1 Fusion ----------- Ø ≠------------ Melting 80 cal g1 °C1 ¯
Water as ice
+ 0 cal g1 °C1 Cooling ---------- Ø ≠-----------Warming 0 cal g1 °C1
where: Condensation = vapour to liquid
Evaporation = liquid to vapour
Fusion = liquid to ice
Melting = ice to liquid
Deposition = condensation and fusion
Sublimation = melting and evaporation
20.6.1 Relative humidity be cooled, without changes in pressure or

water-vapour content, to bring the system to
Relative humidity is the percentage of the water-vapour saturation. At the water-
actual water-vapour pressure in the air com- vapour saturation point, or dew-point/frost
pared with the saturation vapour pressure point, water condenses as liquid water (con-
for that temperature, or the amount of water densation) or deposits as ice (deposition).
the air holds compared with what it can hold. Dew forms if the temperature at which this
Water in the atmosphere serves as a tem- occurs is above 0°C and ice forms when the
perature buffer when it is present, due to its temperature is at or below 0°C. The effect of
energy-holding capacity relative to that of condensation and deposition moderates tem-
the air, its molecular vibrational and rota- perature changes in the system. In the
tional energy and its latent heats of conden- absence of incoming radiation energy, the
sation/evaporation and fusion. During the temperature decrease will be relatively rapid
night, humid air will absorb radiated energy to the dew-point/frost point. Below the dew-
from the earth’s surface and radiate it back. point/frost point, energy is released into the
Low-humidity air allows more energy to be atmosphere from the condensation energy
lost to space. As the air temperature drops given off by water and the rate of tempera-
under humid conditions, the energy stored ture fall is much slower. Growers are encour-
in water is released into the atmosphere at aged when they find they have a high
the rate of 1 calorie g1 of water °C1 and dew-point in the evening before a potential
moderates temperature drop. Under dry air freeze, because they know that the atmos-
conditions, little radiation is absorbed and phere has a high water content and therefore
little heat is given up to the atmosphere, contains much latent energy that will be
resulting in a more rapid and extensive tem- used to moderate temperature fall. In con-
perature drop. trast, if the dew-point is below freezing (i.e.
at the frost point), atmospheric temperature
will drop rapidly to that point, with rela-
20.6.2 Dew-point/frost point tively little energy release. Temperature fall
at the frost point will be moderated by the
The dew-point/frost point of the air is the release of the energy of fusion. But, since the
temperature to which the atmosphere must temperature fall may be rapid to a tempera-
ture below the critical temperature, damage quent energy release would be ineffective in
may occur. This is illustrated in Table 20.3, preventing flower freezing. Freeze B is diffi-
which illustrates two freezes that occur with cult to protect against because only 32 cal-
different amounts of water in the air and, ories (2.67 g of water 12 calories = 32
thus, different dew-points. calories m3) of energy are released from
In freeze A, a high-dew-point freeze, a water per cubic metre in the air above the
total of 689 cal g1 of water in the air would critical temperature. More energy is
be released before the temperature reached released at the ice-formation point of 8°C
the flower’s freezing-point. The most (18°F), but it is released after the flowers
important thing to note is that saturated air have frozen.
in freeze A contains over twice the water The heat exchanges illustrated in the two
and releases about 120 times (5.64 g of freeze situations are approximations only.
water 689 calories = 3886 calories m3) as The situations are actually much more com-
much energy by the time the temperature plex. For instance, dew and ice formation
reaches the critical point than is released in result in changing relative humidities and
freeze B. dew/frost points. These changes result in
In freeze B, a low-dew-point freeze, a changes in subsequent dew/frost formation
total of 12 cal g1 of water in the air would and deposition temperatures so that release
be released before the temperature reached of heat would occur over a range rather than
the critical flower freezing-point and subse- at a point during temperature fall.
Table 20.3. Temperature, water (vapour/dew/ice) and energy relationships in high (A) and low (B) dew-
point freezes.
Temperature Water form Energy release °C1 g1 (calories)
°C ~°F
~ Freeze A Freeze B Freeze A Freeze B
10 50 Vapour Vapour +1 +1
2 36 Dew Vapour +600 +1
1 34 Dew Vapour +1 +1
0 32 Ice Vapour +80 +1
1 30 Ice Vapour 0 +1
8 18 Ice Ice 0 +680
9 16 Ice Ice 0 0
10 14 Ice Ice 0 0
Freeze assumptions:
Critical temperature at full bloom = 2.2°C (28°F)
High-DP freeze A conditions = 10°C DB, 6°C WB, 59% RH, 2°C DP, 5.64 g3 H2O
Low-DP freeze B conditions = 10°C DB, 2.5°C WB, 28% RH, 8°C DP, 2.67 g3 H2O
DB, dry bulb; WB, wet bulb; RH, relative humidity; DP, dew-point.
20.7 Freeze Types sive and dynamic change that frequently

causes damage. Energy reflectants applied to
The Glossary of Weather and Climate of the the south-west side of the tree trunk and scaf-
American Meteorological Society (Geer, 1996) fold limbs prevent this south-west winter
defines a freeze and categories of freezes as: injury. Late-winter freezes may occur when
buds are still relatively hard before apprecia-
The condition that exists when, over a
widespread area, the surface temperature of ble hydration and bud swell has started.
the air remains below freezing 0°C (32°F) for a Spring freezes can occur at increasingly
sufficient time to constitute the characteristic higher temperatures due to acclimatization
feature of the weather. A freeze is a term used accompanying increasing bud hydration and
for the condition when vegetation is injured by metabolic activity. Buds are most susceptible
these low air temperatures, regardless if frost to freezing at full bloom when maximum
were deposited. Freezes may be classified as hydration has opened the flower and tender
light freeze (little destructive damage, except to flower parts are fully exposed to ambient
tender plants); killing freeze (widely destructive
conditions. Critical flower-bud temperatures
to vegetation, effectively terminating the
for apples are given in Table 20.4.
growing season); and hard freeze (staple
vegetation destroyed).
Freezes can also be characterized as 20.8 Monitoring Freezes

advective or radiative. In an advective freeze,
large masses of air, which may have widely Electronic technology makes it possible
varying dew-points, are carried into the today to instrument and monitor various
orchard by wind. The advective freeze may meteorological facets of the orchard with
or may not include clouds. A radiative freeze solid-state devices coupled with computers.
develops when skies are clear, dew-point Temperature-sensing thermocouples should
results in frosts and winds are absent or light. be placed in representative locations verti-
Often a freeze may be a combination advec- cally and horizontally and in the warmest
tive/radiative event. The advective freeze is and coldest areas of the orchard (Plate 20.3).
carried into the orchard by winds; then, after In addition, each block of trees that may
the winds and clouds associated with the respond differently to temperature should be
advection subside, radiative processes occur monitored. These can be interrogated by the
to produce a radiative freeze. computer every few seconds and various
Another classification of freezes depends software packages can provide continuous
on the time of their occurrence. Orchards read-outs of each location’s temperature.
may be damaged in autumn freezes. Autumn National weather services often provide crit-
freezes can be hazardous after conditions that ical information for orchardists. In the USA,
delay tree hardening, such as after autumn the National Weather Service (NWS) fore-
pruning, under fertilization practices that casts minimum temperatures over large
result in growth late in the autumn, or when areas. These temperatures are for standard
tree foliage has been damaged, resulting in thermometers 1.5 m above the ground in
decreased photosynthesis before or during standard NWS instrument shelters. The
the hardening period. Trees may be damaged NWS provides frost warnings when the tem-
in winter freezes when extremely low temperature is predicted to be near 0°C with
peratures occur that exceed the low range for winds less than 16 km h1 (10 m.p.h.),
the species or developmental stage. Autumn frost/freeze warnings when the temperature
and winter freezes may occur in the south- will go below 0°C with winds less than
west side (northern hemisphere) of tree 16 km h1 and freeze warnings when the
trunks and scaffold limbs, due to rapid tem- wind speed is predicted to be above 16 km
perature changes in tissue directly exposed to h1 with temperatures below 0°C. These
the sun during late afternoon. The rapid warnings are estimates for NWS locations
change from an elevated temperature to a and are of limited help to the grower.
freezing temperature at sundown is an exten- Orchardists need specific weather informa-
Table 20.4. Critical temperatures for apple buds during anthesis.

Temperature for 10, 50 and 90% mortality (°C/°F).
Mortality
Bud stagea 10% 50% 90%
Post-dormant
No swelling < 9.4/15.1 < 8.9/15.9 < 17/1.4
Silver tip 9.4/15.1 8.9/15.9 17/1.4
Green tip 7.8/18.0 8.9/15.9 12/10.4
Half-inch green 5.0/23.0 5.6/22.0 9.4/15.1
Tight cluster 2.8/27.0 3.0/27.0 6.1/21.0
First pink 2.2/28.0 2.8/27.0 4.4/24.0
Full pink 2.1/28.0 2.2/28.0 3.9/25.0
First bloom
King 1.7/29.0 2.0/28.4 3.8/25.0
Laterals 2.2/28.0 2.2/28.0 -3.8/25.0
Full bloom 1.7/29.0 2.0/28.4 3.8/25.0
aBud stage refers to average bud stage. Bud populations consist of
some buds ahead and some buds behind the average bud stage. This
population effect produces the critical temperature range. Time is also
a factor; the longer a bud population is at or below the critical
temperature, the more buds will succumb to freezing.
tion for each different area in their orchards sheltered from the sky with inexpensive
and should instrument their plantings to plastic-foam cups (Fig. 20.4). Of course, a
obtain the information required for various short tower such as this is not as good as a
cultural purposes. taller tower, but it can give valuable informa-
Air temperature is the most critical mete- tion to the grower about the air temperature
orological variable to monitor during freeze in the bottom layers of inversions and an
events. Air temperature is usually indica- estimate of the inversion strength. It is also
tive of bud- or flower-tissue temperature, possible to use tethered, instrumented
but it can vary slightly due to radiation bal- helium balloons to monitor inversions under
ance. Therefore, in addition to sheltered calm conditions, but the need to reel the bal-
thermocouples, a grower may want to place loon up and down to obtain temperatures at
small thermocouples closely adjacent to or different elevations is time-consuming.
in buds or flowers to determine their actual Archives of temperature characteristics of
temperatures. inversion events over a few seasons can be
Radiative and advective freezes occur in used to create computer programs that can
almost every orchard every year. To monitor be used to predict radiative-freeze tempera-
radiative freezes, a tower of at least 10 m ture changes through the night as a guide in
should be erected within the block of trees crop protection.
with sensors located on it to measure tem- Another critical meteorological variable
peratures and monitor the inversion that affects temperature change and needs
strength. Two 76 mm (3 in) 6 m (20 ft) sec- to be monitored is the amount of water in
tions of schedule 40 polyvinyl chloride the air or the relative humidity.
(PVC) pipe, coupled together and guy-wired Temperature drop in the orchard during a
for support at 6 and 12 m, can be used suc- radiative freeze is dependent on heat lost
cessfully for a 12 m (40 ft) tower. from the surface during the night. Water in
Thermocouples are placed at 2 m intervals the air modifies temperature changes; there-
on the tower. Thermocouples need to be fore, monitoring water in the atmosphere is
from the heat of condensation (stored or

latent heat) is released. This heat, when
added to the atmosphere, decreases the rate
of temperature fall significantly. Above the
dew-point, temperature fall is relatively
rapid, but, after dew formation begins, the
temperature fall is relatively slow. If the
dew-point is significantly above the critical
temperature, there is little danger of a criti-
cal freeze, but, if the dew-point is near or
below the critical killing temperature of
flower buds, the temperature will drop
rapidly into the critical area and protection
will be needed.
Significant advantages accrue to growers
who educate themselves in meteorology and
use historical records and current tempera-
ture and dew-point monitoring to estimate
critical temperature probabilities to predict
potential freezes in their blocks of trees.
There are custom forecasts available, but,
even when they are accurate, they are not as
good as continuous monitoring of the meteo-
rological variables that affect temperature
fall in a specific orchard during potentially
hazardous freeze nights.
Fig. 20.4. Sheltered (plastic-foam cups cut in half)

thermocouples on a 4 m tower. 20.9 Methods for Freeze Protection
20.9.1 Heating
very important, because it will reflect the
heat energy available to moderate tempera- Orchards have been heated during freezes
ture fall. Clouds, for instance, create a ther- with anything that would burn. Prunings,
mal blanket for the orchard. When clouds firewood, tyres, coal, oil, diesel, charcoal,
are present, they create a greenhouse effect coke, paraffin, solid petroleum, liquid
and contain energy in the atmosphere. Very petroleum, propane and many other materi-
thin clouds as high as 10,000 m can create als have been used. Systems have pro-
this effect. gressed from open fires on the orchard floor,
When clouds are not present, humidity to metal pots, to baskets in trees, to return-
needs to be monitored to provide a reason- stack heaters, to specialized high-radiative-
able estimate of the rate of temperature fall. output heaters fed by underground fuel
A psychrometer (a hygrometer consisting of lines connected to underground tanks.
two similar thermometers, with the bulb of Individual heaters (up to 100 return-stack
one kept wet, so that the cooling that results heaters and 150 pressurized-propane
from evaporation makes it register a lower heaters ha1) can be located throughout the
temperature than the dry one, and with the orchard to obtain the desired heat distribu-
difference between the readings constituting tion (Plate 20.4). Many small heaters spread
a measure of the wetness of the atmosphere) throughout the orchard to allow uniform
can be used to determine the dew-point. The heating under various air flows and site
dew-point is important in the rate of characteristics are much better at maintain-
temperature fall. At the dew-point, energy ing temperatures above the critical freezing-
point than just a few larger heaters. Large 20.9.2 Overhead irrigation
heaters can release enough heat to create a
chimney effect through the top of the inver- The temperature of an ice/water coating on
sion, with heat losses near the ground. trees remains very close to 0°C. Ice build-up
However, large heaters can be used under results in much more ice than water on the
wind machines in a situation where the ris- tree, with the water in a thin surface film.
ing heat plume is distributed horizontally Water is added rapidly enough with over-
through the orchard by the wind machine. head sprinklers to form small clear icicles,
Usually heaters are more concentrated on which remain covered with a thin water film.
the orchard borders, especially on the wind- Icicles may form in arcs due to the changing
ward side. The pipeline-fed systems have load on the limbs. Ice with entrapped air,
many fixed heaters per acre that are not which appears white, indicates that water is
readily moved from place to place so that not being added at a sufficient rate to main-
the optimum arrangement of heaters for the tain a constant ice/water coating. Sprinkler
most characteristic freeze conditions must heads should be of a design that will con-
be made at the time of installation. Pumps, tinue to rotate with ice build-up. Gravity
filters, pressure controls, automatic lighting sprinkler systems are efficient, but pumped
devices, pilot lights, specialized nozzles and systems in which the pressure can be
computer control are components that char- increased are better, since they can provide
acterize the most elaborate systems. The variable water-application rates. Ice/water
cost of return-stack oil heaters is around encasement of apple trees before and at full
US$2600 ha1. Pressurized propane bloom has been used successfully for many
heaters, including some to heat the storage years. It is not used after full bloom because
tank, along with the service equipment encasement of foliation associated with
mentioned, may cost from US$6200 to shoot growth results in excessive ice build-
US$10,000 ha1. up and may damage tree structure. Smaller
Most commercial heaters and home-made trees trained to central-leader systems, such
heaters from surplus shell casings emit the as the Pacific Northwest central leader,
majority of their heat as convective heat and Hytec, slender-spindle and vertical-axis sys-
thus they are very effective under inversion tems, easily withstand the load of ice encase-
conditions. However, these heaters do not ment. Open-centre trees do not tolerate ice
perform as well under advective conditions loading as effectively as central-leader trees.
as heaters having a high radiative output. Significant quantities of water are used
Some heaters – for instance, coke heaters that for ice/water-encasement freeze protection
produce ~40% of their heat as radiant heat – (Fig. 20.5). Soils subjected to water saturation
have a greater effect on bud temperature during this procedure should be adequately
under advective conditions, since radiation drained so that root function will not be
is not disturbed by winds. impaired due to water accumulation or low
The combination of heaters and wind soil temperatures. Irrigation systems used
machines, which allows mixing of an exist- for ice encasement for freeze protection
ing inversion with heat additions within the should be installed following standard irri-
inversion, has been successful. Increases in gation design principles, but they should be
fossil-fuel costs have made most systems over-designed to allow sufficient rates of
obsolete. However, wind machines and water application to protect against freezes
heaters can be used for citrus-orchard protec- down to about 8°C (18°F). The system
tion, where the goal is not only to save the should be capable of applying up to 1.2 cm
current fruit crop but also the orchard itself. h1 for freezes near the critical temperature
The combination can also be used in blocks of the green-tip bud stage of –8°C (18°F) to
of high-density, high-value deciduous the critical temperature at full bloom of –2°C
orchards to save the trees during severe win- (28°F). Sufficient water must be readily avail-
ter freezes or to ensure a crop for specific able to continue the sprinkling until all ice
lucrative retail markets. has melted.
Fig. 20.5. Ice build-up from sprinkler overhead irrigation.
During a freeze, the ice/water-encase- 20.9.3 Under-tree sprinkling

ment sprinkler system should be started at a
temperature several degrees above the criti- Several apple growers have reported success
cal to avoid possible killing temperatures with under-tree sprinkling for freeze protec-
due to evaporative cooling on system start- tion. Large experiments in Utah (USA), using
up. Since the heat of fusion of water on elaborate statistical designs, in a tart-cherry
freezing is 80 cal g1 and the heat of vapor- orchard in the spring seasons of 3 different
ization associated with evaporation is 600 years gave no positive results (Plate 20.5). Of
cal g1, the evaporation of 1 g of water course, the high mountain desert with its low
absorbs the heat equivalent to the freezing humidity would be expected to result in high
of 7.5 g. Winds and the relative humidity of evaporative cooling and, indeed, the only sig-
the air influence the evaporation rate, but nificant temperature differences were nega-
relative humidity is modified greatly as tive. Others, however, have reportedly used
soon as sprinkling is initiated. Therefore, the under-tree sprinkling successfully for freeze
forecast minimum temperature and wind protection. Orchardists in the Casa Grande
speed are critical in determining the rate of area of Mexico use 21°C (70°F) well water to
water application. If the forecast tempera- protect their orchards and report success. It
ture minimum is below the protection has also been reported that orchardists in
capacity designed into the system, the sys- Washington State (USA) have used heated
tem must not be started because severe water to provide under-tree sprinkling for
damage can result. Wind speeds of 3 m s1 freeze protection in their orchards.
require more than twice the amount of
water for protection as that needed for
winds of 0.5 m s1. At low wind speeds, as 20.9.4 Wind machines
temperatures drop from 2 to 8°C (28 to
18°F) – about the practical limit for ice The climatology of a site must be analysed to
encasement protection – water requirements determine the incidence of advective and
increase approximately 0.1 cm h1 °C1. radiative freezes because wind machines are
Ice/water-encasement-use decisions need only useful under radiative freeze conditions
to incorporate the critical temperature, the when a significant inversion exists. Other
forecast minimum temperature in the types of protection, such as coverings or
orchard, the wind speed, the dew-point and bloom delay, must be provided during
the water-delivery capacity of the system. advective freezes.
During an inversion freeze, the air above ardous areas, flying at low altitudes and the
the orchard is warmer than the air near the necessity of refuelling and service stops are
ground (Fig. 20.3c). Radiation from the sur- problematic in obtaining the desired contin-
face, conduction of heat from the air near uous inversion mixing.
the surface to the surface and the absence of Wind machines should be located in the
convective energy transfer result in the typi- centre of the area to be protected or slightly
cal inverse-temperature curve in relation to to the upwind side. Fans rotate through
the curve during the daytime. Inversions 360° in 3–4 min. Most machines have 4 m
develop on clear, calm nights, which allow diameter propeller blades mounted at a
radiational and conductive-to-radiational very slight angle at the top of 10 m towers.
heat losses from the surface. Inversions are Single-propeller machines of about 1.86
responsible for fogs, valley clouds and pol- 105 W s1 can protect about 4 ha, and
lutant trapping in the lowest layers of the double-propeller machines can protect
atmosphere. The temperature stratification about 6 ha. Machines should be serviced
that develops during an inversion is diffi- regularly, with tuning, fuelling, charging,
cult to modify. Significant power – approxi- lubricating and tachometer checks and
mately 1.86 104 W s1 ha1 – is necessary mechanical checks of all moving parts and
to modify an inversion with wind machines their fasteners. The energy consumption of
sufficiently to provide freeze protection. wind machines is low relative to that of
Temperature modification can equal about heaters. Machines need to be started before
one-quarter of the inversion-temperature they are needed and idled until they are
difference or one-half of the temperature used. Wind machines with supplementary
difference between 2 and 20 m. Once the heaters are more economical than heaters
protection has been established, the only when additional heat is needed. Wind-
machine must keep working or the inver- machine/heater synergy ranges from 20 to
sion will re-establish. 30%. One study found that the combined
Wind machines should be started before
response of a wind machine and 20 heaters
the temperature approaches critical, since it
ha1 is about equal to about 55 heaters ha1
is easier to maintain a temperature than to
alone. Machines of 1.86 105 W s1 on
produce an increase in temperature.
12 m towers supplemented with about 93
Inversions will not be present when natu-
heaters ha1 evenly spaced throughout the
rally occurring winds mix the air, when
area worked very well in another freeze. Of
clouds are present to reradiate energy back
course, heater density and placement
to the surface or when cold air can drain
should counteract colder upwind borders
away from a site. Wind-machine mixing of
and corners.
the air above the orchard with the air in the
orchard tends to equilibrate the energy in the
system making orchard temperatures higher
20.9.5 Bloom delay
(Plate 20.6). Wind-machine air movement is
proportional to the propeller r.p.m. High fan
20.9.5.1 Genetic control
speeds are necessary for the desired temper-
ature modifications. Time of apple flowering is influenced by
Helicopters – the larger the better – are the climate and the genotype, with full
effective in mixing inversions. The pilot can bloom dates varying by as much as 6
monitor the inversion and operate at the weeks. Most cultivars, however, bloom
optimum altitude. Thermometers that trig- within a 3-week period. The time of leaf-
ger small lights at temperatures slightly bud burst is correlated with the date of
above the critical temperature can be placed flowering and cultivars can be selected for
throughout the orchard and the helicopter late flowering on the basis of leaf-bud
can move so as to extinguish the lights burst. Seeds of late-flowering genotypes
immediately after they turn on. Scheduling have higher chilling requirements and ger-
helicopters, flying at night, flying over haz- minate over a much longer period than
seeds of early-flowering genotypes. Commercial bloom-delay systems have used

Dormancy is a complex polygenic trait. impact-type sprinklers, which usually
Therefore, expensive, time-consuming deliver too much water and may cause soil
breeding has been limited to fruit quality waterlogging and exacerbate diseases.
and resistance to disease, with little atten- Water-application systems utilizing umbrella
tion to hardiness or bloom date. and misting nozzles, controlled by leaf-wet-
ness gauges, have been more successful in
producing delays without over-application
20.9.5.2 Bioregulators
of water, but they are more expensive. All
Bioregulators, including the plant growth systems require automatic water flow and
hormones ethylene, auxin, gibberellin and draining equipment. Calcium carbonate
abscisic acid, have been used to delay bloom build-up on trees in some studies did not
of several species of orchard trees. While have an adverse effect on subsequent tree
they do give some bloom delay, in general, growth or production.
they have too many side-effects to be useful. Approximately 25% of the thermal energy
Oils, 1,2-dihydro-3,6-pyridazinedione (maleic required for apple flower-bud development
hydrazide), butanedioic acid mono(2,2- from the end of endodormancy to full bloom
dimethylhydrazide) (daminozide), 2-chloro- is utilized to develop to bud swell, the first
N,N,N-trimethylethanaminium chloride stage of observable phenological develop-
(chlormequat), and aminoethoxyvinylglycine ment. Another 25% is received by the time of
(AVG) have also proved to be ineffective or the tight-cluster stage of development. When
contraindicated or to delay apple bloom for evaporative cooling for bloom delay is
only a limited time. applied correctly within these phenological
limits (end of endodormancy to tight-cluster
stage), it will be successful and will avoid
20.9.5.3 Evaporative cooling
most of the complications – fire blight, apple
Evaporative cooling has been used to delay scab, waterlogging, delayed root growth, fer-
bloom of apple for as long as 17 days. The tilizer leaching and hydration of flower
technique requires determination of the end buds, which results in decreased hardiness –
of endodormancy and is facilitated by moni- connected with water applications nearer
toring of anthesis phenology. Water applica- full bloom.
tions to induce evaporative cooling, which It is important to apply evaporative cool-
are more effective in arid climates with low ing during the time when the tree is still rela-
humidities that facilitate evaporation, should tively dormant and there is time for
begin shortly after endodormancy ends. To water-saturated soils to drain before root
initiate cooling, a thin coat of water should growth begins. In one study, evaporative
be applied to the tree when temperatures in cooling was applied for 348 h over a 40-day
the orchard are several degrees warmer than period to produce a 17-day bloom delay;
the approximate 5°C (41°F) threshold of sprinkling for a subsequent additional 238 h
apple-tree growth. Evaporation of the water produced only an additional 3-day delay.
absorbs thermal energy from the tree and the Bloom delay in short-growing-season pro-
environment, resulting in mid-afternoon duction areas produces mature apples with
temperature differences of as much as 20°C the same size and colour but with lower sol-
(36°F) between sprinkled trees and their con- uble solids than non-delayed fruit, but the
trols. In dry areas, evaporative cooling is difference in harvest time is only a fraction of
more than 90% efficient in reducing fruit-bud the delay in bloom.
temperatures to the wet-bulb temperature.
Significant bloom delay can be obtained in
20.9.5.4 Energy reflectants
these areas (Plate 20.7).
Almost all of the applied water should be Energy reflectants can be applied to buds
allowed to evaporate to produce the cooling and limbs of trees during the winter to pro-
effect before more water is applied. duce a bloom delay of several days. The tem-
Table 20.5. Use of freeze-protection methods.
Freeze type
Method Advective Radiative Advective/Radiative
Heaters No Yes No/Yes

Wind machines No Yes No/Yes
Ice encasement No Yes No/Yes
Under-tree sprinkling No Yesa No/Yesa
Bloom delay Yes Yes Yes/Yes
Site selection Yes Yes Yes/Yes
aUnder-tree sprinkling does not work in some areas.
perature of the tree remains lower than that 20.10 Summary

of control trees due to reflection rather than
absorption of radiant energy, so that devel- Freeze-protection-method use is summa-
opment is slower. rized in Table 20.5. All freeze-protection
methods listed have been used successfully
during radiative freezes, but only site selec-
20.9.5.5 Shading
tion and bloom delay are effective in avoid-
Trees that are shaded during anthesis so that ing freeze losses due to advective freezes.
their temperatures are lower than ambient Protection during advective/radiative
will bloom several days later than those freezes depends on the severity of each com-
growing in direct sunlight. ponent. If the advection is relatively mild,
with cloud cover and humid air to moderate
temperature drop, and the crop survives the
20.9.5.6 Trunk refrigeration
advection freeze, the methods that are suc-
Trunk refrigeration has been used to delay cessful during radiative freezes may save the
bloom of apple and peach trees. However, crop. However, if the advection is severe,
expensive refrigeration apparatus, distribu- crops will be lost. Therefore, selection of a
tion pumps, lines, insulation and specialized freeze-free site is by far the best solution. All
equipment to regulate water stress in the tree other solutions are more expensive and
make the method costly and impractical. make the enterprise less profitable.
Reference and Further Reading
Ahrens, C.D. (1998) Essentials of Meteorology: an Invitation to the Atmosphere, 2nd edn. Wadsworth,
Belmont, California, 443 pp.
Anderson, J.L. and Seeley, S.D. (1993) Bloom delay in deciduous fruits. Horticultural Reviews 15, 97–144.
Barfield, B.J. and Gerber, J.F. (1979) Modification of the Aerial Environment of Plants. Monograph, American
Society of Agricultural Engineers, St Joseph, Michigan, 538 pp.
Geer, I.W. (1996) Glossary of Weather and Climate with Related Oceanic and Hydrologic Terms. American
Meteorological Society, Boston, Massachusetts, 272 pp.
Perry, K.B. (1998) Basics of frost and freeze protection for horticultural crops. HortTechnology 8, 10–15.
Rieger, M. (1989) Freeze protection for horticultural crops. Horticultural Reviews 11, 45–109.
Apples - Ch.21 21/3/03 3:03 pm Page 539
21 Integrated Fruit Production for Apples –

Principles and Guidelines
Jesús Avilla1 and Helmut Riedl2

1Centro UdL-IRTA de R+D de Lleida, University of Lleida, Lleida, Spain;
2Mid-Columbia Agricultural Research and Extension Center,
Oregon State University, Oregon, USA
21.1 The Concept of Integrated Fruit Production (IFP) 539

21.2 The Principles of IP 540
21.3 Integrated Production for Apples 541
21.4 IFP Implementation/Adoption 543
21.5 Case-study 1: Integrated Production for Apples in South Tyrol (Italy) 544
21.6 Case-study 2: Integrated Production for Apples in Patagonia (Argentina) 545
21.7 Case-study 3: Integrated Production for Apples in New Zealand 546
21.8 Case-study 4: Integrated Production for Apples in Northern Oregon (USA) 547
21.1 The Concept of Integrated Fruit (http://www.admin.ch/sar/faw/iobc) is

Production (IFP) responsible for developing and administer-
ing IP guidelines, while several working
The development of integrated production groups provide crop-specific technical infor-
(IP) has been the result of the collaborative mation. The ISHS was established in 1959,
effort of many scientists, mainly during with the aim of promoting and encouraging
the last 25 years, and of two leading research in all branches of horticulture.
organizations: the West Palearctic Regional The term ‘integrated production’ was
Section of the International Organization coined at a meeting organized by IOBC/
for Biological and Integrated Control of WPRS, held at Ovrannaz, Switzerland, in
Noxious Animals and Plants (IOBC/WPRS) 1976. As a result of this meeting,
(http://www.iobc-wprs.org) and the Inter- IOBC/WPRS published a document that
national Society for Horticultural Science is considered the starting-point for IP in
(ISHS) (http://www.ishs.org). IOBC/WPRS Europe (Steiner, 1977). Since then,
is one of six regional sections of the IOBC; IOBC/WPRS has been the leading organiza-
it is an independent, non-profit, scientific tion in promoting research on and extension
organization established in 1956 to promote of IP and has published several guidelines,
environmentally safe methods of pest (ani- both general and crop-specific, some of them
mals, pathogens and weeds) control in plant in collaboration with the ISHS. The guide-
protection and, later, to promote the lines can be downloaded from the
development and adoption of IP methods. IOBC/WPRS IP Commission internet home-
The IP Commission of the IOBC/WPRS page mentioned above.
Apples - Ch.21 21/3/03 3:03 pm Page 540
540 J. Avilla and H. Riedl
IOBC/WPRS has defined IP as a ‘farming basis for the planning and realization of all
system that produces high-quality food and farm activities, particularly those with poten-
other products by using natural resources and tial ecological impact. They are the visible
regulating mechanisms to replace polluting expressions of the holistic concept and pro-
inputs and to secure sustainable farming’ vide both a natural resource and a manage-
(Boller et al., 1999). Within the framework of ment component. The entire farm is then the
this general definition, IFP is defined as ‘the basic unit for IP implementation, as IP
economical production of high quality fruit, applied on isolated individual areas is not
giving priority to ecologically safer methods, compatible with achieving a holistic
minimizing the undesirable side effects and approach. Important strategies, such as
use of agrochemicals, to enhance the safe- achieving balanced nutrient cycles and hav-
guards to the environment and human health’ ing an optimum allocation of farm machin-
(Cross and Dickler, 1994). These definitions ery, only become meaningful if considered
have been widely accepted worldwide and across the entire property. Some of the IP
are part of the national regulations of several principles (such as the maintenance of stable
countries, such as Switzerland. agroecosystems and the support of biological
Unlike organic farming, IFP does not seek diversity) should be applied on an even
to eliminate the use of agricultural chemicals wider scale and, in fact, are more easily
but rather to reduce (or even eliminate in applied across large areas. For that reason,
some cases) production inputs with high IOBC/WPRS focuses on the importance of
environmental impacts, such as broad-spec- growers’ associations to implement IP pro-
trum pesticides or fertilizers, and to favour grammes.
safer alternatives (Sansavini, 1990, 1997; IPM is a key component of IP. Numerous
Plate 21.1). case-studies have shown that the adoption of
a sound IPM programme is often the first
step in implementing IP practices (see, for
21.2 The Principles of IP example, Case-study 1). To avoid misunder-
standing, the widely recognized definition of
The principles of IP, according to IPM is:
IOBC/WPRS, are shown in Table 21.1 (Boller
A pest management system that, in the context
et al., 1999). IP represents an ecological
of the associated environment and the
approach to crop production and relies on population dynamics of the pest species,
ecosystem management and on the preserva- utilizes all suitable techniques and methods in
tion of natural resources. It is a holistic con- as compatible a manner as possible and
cept and is not merely a combination of maintains the pest populations at levels below
several elements, such as integrated pest those causing economic injury.
management (IPM), together with some (FAO Panel of Experts on Integrated
agronomic measures. Agrosystems are the Pest Control, 1967)
Table 21.1. IP principles according to IOBC/WPRS (from Boller et al., 1999).
1. IP is applied only holistically

2. The external costs and the undesirable impacts of agriculture are minimized
3. The entire farm is the unit of implementation of IP
4. The farmers’ knowledge of IP must be regularly updated
5. Stable agroecosystems are to be maintained as key components of IP
6. Nutrient cycles are to be balanced and losses minimized
7. The intrinsic soil fertility is to be preserved and improved
8. Integrated pest management is the basis for decision making in crop protection
9. Biological diversity must be supported
10. The quality of the final products must be evaluated using the ecological parameters of the production
system, as well as by the usual external and internal quality parameters
Apples - Ch.21 21/3/03 3:03 pm Page 541
Integrated Fruit Production for Apples 541
IP and IFP are, then, much broader concepts processing and handling of products is a
than IPM or integrated plant protection (IPP) prerequisite for the IP label.
and are derived from the expansion of the The principles of IPP and IP, as well as the
principles of IPM to all field practices description of the evolutionary steps from
(Sansavini, 1990). The emphasis of plant pro- chemical control to IP, have provided impor-
tection in the context of sustainable agricul- tant orientation marks for the development
ture is placed on preventive measures of a sustainable approach in agriculture
(‘indirect plant protection’), which must be during the last two decades (Boller et al.,
utilized to the fullest extent before direct 1998).
plant-protection measures (i.e. control mea-
sures) are applied (Boller et al., 1998).
Decisions about the necessity to apply con- 21.3 Integrated Production for Apples
trol measures must rely on the most
advanced tools, such as prognostic methods The third edition of the IOBC/WPRS
and scientifically verified thresholds. The guidelines for the IP of apples was pub-
application of direct plant-protection meth- lished in 2002 (Cross, 2002) and the full
ods is only used if economically unaccept- text is available at the IOBC/WPRS IP
able losses cannot be prevented by indirect Commission internet site, mentioned above.
plant-protection methods. The key items covered in the document and
Biological diversity includes diversity at discussed below are summarized in Table
the genetic, species and ecosystem level. It is 21.2.
the backbone of ecosystem stability, natural At least two ecological options for the
regulation factors and landscape quality. active enhancement of biological diversity
Replacement of pesticides by natural regula- are required, such as the use of nesting
tion factors cannot adequately be achieved boxes and/or perches for predatory birds,
without adequate biological diversity. The refuges for predators, host plants for benefi-
conservation and enhancement of biological cials and new wildlife habitats. The increase
diversity is therefore an important element in biodiversity is more easily achieved in
of IP (Boller et al., 1999). IOBC/WPRS IP more or less permanent agroecosystems,
guidelines include, as mandatory, the exis- such as apple orchards, mainly in cool cli-
tence of an ecological compensation area, mates, where a permanent green cover can
which must cover at least 5% of the total be maintained all year round (Boller et al.,
farm area (excluding any area of forest). The 1998).
ecological compensation area includes areas For new orchards, site selection, root-
with no input of fertilizers and pesticides, stocks, cultivars and planting systems must
such as hedges, natural areas and field be selected and harmonized so that regular
boundaries. Special cases might be individu- yields of quality fruit, and hence economic
ally owned orchards and vineyards within success, can be expected with the minimum
larger complexes of crop production. In this use of both agrochemicals and other envi-
case, IOBC/WPRS recommends that the IP ronmentally hazardous practices. Chemical
organization identifies the 5% area of the soil sterilization is not permitted. The culti-
entire area owned by its members (Boller et var chosen must offer good prospects for
al., 1998). economic success with minimal use of agro-
The quality of products obtained under IP chemicals. For example, ‘Golden Delicious’
systems must be measured not only in terms must not be planted on sites prone to russet-
of external and internal quality (which must ing or ‘Jonagold’ on sites unfavourable for
be as high as under conventional produc- fruit colouring and firmness. Cultivars resis-
tion), but also in terms of impacts on the tant or tolerant to diseases and/or pests are
environment and on human health. Hence preferred. Planting material should be of
certification testifying to the achievements of high quality and certified as being virus-
the producer and defining the requirements free.
that have been met during the storage, Soil must be sampled and chemically
Apples - Ch.21 21/3/03 3:03 pm Page 542
Table 21.2. Items covered by the IOBC/WPRS–ISHS guidelines: integrated production of apples (from
Cross, 2002).
1. Definition of integrated production of pome fruits

2. Professionally trained, environmentally and safety-conscious growers
3. Conserving the orchard environment
4. Site, rootstocks, cultivars and planting systems for new orchards
5. Soil management and tree nutrition
6. Alleyways and weed-free strip
7. Irrigation
8. Tree training and management
9. Fruit management
10. Integrated plant protection
11. Efficient and safe spray-application methods
12. Harvesting, storage and fruit quality
13. Postharvest chemical treatments
analysed prior to planting. Regional or a maximum width for the weed-free strip
national guidelines must set out a clear and/or percentage of the soil surface that
method by which fertilizer requirements are may be weed-free.
determined and define both sampling and Where possible, in established cropping
analytical procedures and the rules on which orchards with excessively vigorous growth,
decisions are to be made. It is recommended the use of herbicides must not be permitted.
that N-min tests be used. The N-min test To avoid undue competition for moisture
determines the existing share of mineral and nutrients, a weed-free strip should be
nitrogen (nitrate and ammonium) in the soil. maintained by mulching or by covering the
On the basis of humus content and soil type, soil surface or through the use of mechanical
the nitrogen replacement value of the tree cultivation. It is recommended that, where
row is estimated and, finally, the nitrogen possible, ground cover is allowed to develop
fertilization is calculated. The maximum in the weed-free strip at times of year (such
nitrogen input (expressed in kg ha1 year1) as in winter) when soil moisture is adequate.
and the timing and methods of application Herbicides must not be used to achieve over-
must be defined and managed in order to all bare soil and the avoidance of the use of
minimize leaching. The same principles selective broad-leaved weed herbicides in
apply to all other major nutrients with high the alleyways is recommended.
polluting potential. Trees must be trained and pruned to
The aims of ground-cover management are: achieve a manageable uniform size and a
balance between growth and regular yields
● to maintain plant-species diversity in the
and to allow good penetration of light
orchard, thus fostering ecological stability;
and spray chemicals to the tree centre. The
● to minimize the use of herbicides (avoid-
use of non-naturally occurring, synthetic,
ing residual chemicals completely);
plant growth regulators is not permitted.
● to avoid soil erosion and compaction in
Excessive growth should be controlled by
alleyways, without detriment to yield and
cultural measures, including the use of
with minimum inputs of fertilizers and
reduced fertilizer and irrigation supply, sum-
irrigation water.
mer pruning and approaches to achieve
Bare-soil management over the entire greater blossom set.
orchard is not permitted. Alleyways must be Priority must be given to natural, cultural,
of grass and/or herbs and be of adequate biological, genetic and biotechnical methods
width to accommodate easily a vehicle of pest, disease and weed control, and the
passing along the rows. Non-competitive use of agrochemicals must be minimized.
grass/herb mixtures are recommended. Plant-protection products may only be used
Regional or national guidelines must specify when justified and the most selective, least
Apples - Ch.21 21/3/03 3:03 pm Page 543
toxic, least persistent products, which are as Comité Française pour la Valorisation de la
safe as possible to humans and the en- Production Fruitière Intégrée (COVAPI) in
vironment, must be selected (Plate 21.2). France, began to embrace IFP practices. In
Populations of key natural enemies (such as 1989, the Arbeitsgruppe für den integrierten
phytoseiid mites on apple or anthocorid Obstbau in Südtirol (AGRIOS), an IFP orga-
predators on pear) must be preserved. At nization in the apple-growing area of north-
least two key natural enemies in each crop ern Italy, published the first comprehensive
must be identified in national/regional IFP guidelines (see Case-study 1). Fruit
guidelines. This means plant-protection grown according to the guidelines is certified
products toxic to them may not be used. and sold with a special label that identifies it
Where phytoseiid predators are absent from as coming from IP. The publication of the
apple orchards, they should be introduced AGRIOS guidelines was an important event,
where necessary. There are some excellent since it stimulated similar efforts in other
books on IPM of apple pests (Beers et al., European fruit-growing areas. In 1991, work-
1993; Statewide Integrated Pest Management ing groups of IOBC/WPRS and the ISHS
Project, University of California, 1999). jointly developed IP guidelines for pome
The following criteria should be taken fruits. The goal was to harmonize regional
into account in the selection of pesticides for guidelines and set European standards for
use in IP programmes: IFP. Many European fruit-growing areas have
now adopted IFP programmes, which gener-
● toxicity to man;
ally follow the principles set forth by the
● toxicity to key natural enemies;
IOBC/WPRS guidelines for pome fruits.
● toxicity to other natural organisms;
Major apple-growing countries around the
● pollution of ground- and surface water;
world have followed the European example
● possibility of provoking an increase in
and have developed their own IFP pro-
pest populations;
grammes, including South Africa, Argentina
● selectivity;
(see Case-study 2), Brazil, Chile, Uruguay and
● persistence;
New Zealand (see Case-study 3). Sustainable-
● incomplete information;
agriculture research and education pro-
● necessity of use.
grammes similar to IFP are also under way in
Postharvest treatment with synthetic, non- several fruit-growing areas in the USA, such
naturally occurring antioxidants for control as Oregon (see Case-study 4), but they still
of superficial scald and other disorders is not lack certification and marketing components.
permitted. In order to minimize the use of Several reasons may explain the rapid
fungicide sprays shortly before harvest for increase in the number of IFP programmes
control of storage diseases, postharvest that followed the publication of the first
fungicide treatment of fruit is permitted AGRIOS guidelines in 1989:
where several conditions have been fulfilled
1. The AGRIOS programme underlined, for
(Cross, 2002).
the first time, the economic benefits of IFP in
terms of production costs, marketing oppor-
tunities and consumer safety (Dickler and
21.4 IFP Implementation/Adoption Schaefermeyer, 1993). IP fruit does not gener-
ally get a higher price, but several large-scale
The development of IFP in Europe was European produce distributors prefer it to
closely linked to the early work on apple IPM conventionally produced fruit, since the label
in the 1960s (Dickler and Schaefermeyer, ‘from integrated production’ adds value (i.e.
1993). However, the adoption of IFP practices ecological quality) in the eyes of distributors
by growers was slow and took many years. as well as consumers. Although both organic
In the 1970s, the first grower organizations, and IP fruit could meet the requirements for
such as the Groupement des Arboriculteurs ecological quality, only IFP is expected to
Lémaniques Pratiquant les Techniques meet these objectives at the commercial level
Intégrées (GALTI) in Switzerland and the (Sansavini, 1990; Boller et al., 1998).
Apples - Ch.21 21/3/03 3:03 pm Page 544
2. The interest of consumers in a ‘safe and fruit-growing areas in Europe. It stretches for
healthful food supply’ is increasing. This about 110 km along the Etsch valley, which is
trend began in the 1980s and is continuing 3 km wide on average and is flanked by high
(Boller et al., 1998). mountain ranges. The altitude varies from
3. The agricultural policy of the European 300 to 1000 m above sea level. The soils are
Union (EU) actively promotes the adoption mostly light-textured, permeable, sandy
of IFP practices. Specific EU regulations for loam soils with a pH of 5.5–7.0 and a fairly
IP practices are still lacking at this time. balanced nutrient content. The average rain-
However, the EU financially supports, fall varies from 600 to 800 mm year1. The
through subsidies, grower organizations minimum winter temperature may reach
as well as individual growers who have 10°C, exceptionally 15°C, and the maxi-
operational programmes in place whose objec- mum summer temperature may reach 30°C.
tives include the ‘promotion of Integrated The fruit farms are small, with an average
Production or other methods which respect size of 3–4 ha. The orchards are scattered
the environment’ (EU Directive 2200/96). It across the slopes and the floor of the Etsch
is expected that the EU will eventually adopt river valley. At present, there are about 8000
uniform IFP standards that will apply to all orchard owners. The area devoted to apple
member countries. production in South Tyrol is 17,600 ha.
In 2001 the total pip-fruit production was
Apple acreage under IFP programmes has
940,000 t.
been steadily increasing in Europe.
The history of apple (and pear) produc-
According to a survey conducted in 1994, 35%
tion in South Tyrol shows the progression
of the apple production area was under IFP or from conventional agriculture to IFP through
similar systems. This represented a 40% IPM. The occurrence of several cases of pest
increase in IFP acreage since 1991 (Cross et al., resistance (European red mite in 1964 and
1996). The same survey found a considerable 1982, leaf-miners in 1969 and 1988, pear
drop in pesticide use in orchards under IFP psylla in 1973) led the South Tyrolean
programmes. A more recent survey, conducted Advisory Service to implement IPM pro-
in 1997, showed that the adoption of IFP was grammes (1977–1987) through the organisa-
considerably higher in central and northern tion and supervision of 15 grower working
Europe (E. Dickler and E. Olivella, Spain, 1999, groups. A reduction of the number of insecti-
personal communication). For instance, in cide and acaricide applications and an
Switzerland, where more than 70% of apple increase of biological control (woolly apple
production is under IFP, it has become difficult aphid and European red mite) were already
to market fruit if it does not have an IFP label. occurring at this time.
The same is true for New Zealand, where all As a result of the initiative of the institu-
fruit destined for export must have an tions and organisations involved in fruit
approved IFP label. It is expected that IFP or production in South Tyrol, the Working
similar production systems will become the Group for Integrated Production in South
standard for growing apples and other tree Tyrol (AGRIOS) was formed in 1988. The
fruits in Europe and other fruit-growing areas. first edition of the AGRIOS guidelines was a
milestone in European IP, and it was pub-
lished in 1989 under the supervision of Dr
21.5 Case-study 1: Integrated Hermann Oberhofer. At present, AGRIOS
Production for Apples in activities include the elaboration and publi-
South Tyrol (Italy)1 cation of guidelines for fruit production and
storage; the adjustment of the IFP pro-
The South Tyrol, also called Alto Adige gramme to meet production and marketing
(northern Italy) is one of the biggest pome- requirements; the administration of the
1 Source:Wierer, P., AGRIOS. Workgroup for Integrated Fruit Production in South Tyrol, and Waldner,
W., Südtiroler Beratungsring für Obst und Weinbau, http://www.agrios.it
Apples - Ch.21 21/3/03 3:03 pm Page 545
participating producers and packing- the professionalism of the growers has

houses; the preparation, publication and increased, the image of South Tyrol produc-
distribution of the AGRIOS newsletter (c. 12 tion is much better and new markets have
a year, 6500 copies per newsletter); and the been opened.
control of the participating farms and pack-
ing-houses. AGRIOS guidelines follow the
conceptual framework of IOBC/WPRS. 21.6 Case-study 2: Integrated
They give a special emphasis to the ecologi-
Production for Apples in Patagonia
cal measures that have positive impacts on
(Argentina)2
the orchard and the environment and to the
use of biological and biotechnical pest-con-
The Alto Valle del Río Negro (Upper Valley
trol measures. A list of ecological measures
of the Negro River) is an Argentinean region
is given and the grower is encouraged to
whose main economic activity is fruit pro-
apply as many of them as possible each year
of production. Some of the ecological duction. Its production of apples and pears
options are: green cover all year round; represents about 80% of the national produc-
hanging up of nesting boxes; creation of tion and most of it is dedicated to export.
refuges for weasels, hedgehogs and other The climate is temperate, continental and
animals; introduction of phytoseiid mites; arid, making irrigation necessary.
planting of hedges and bushes in the edges The development of the IP programme
of the orchards; and installation of perches for apples and pears began in 1994, with the
to attract birds of prey (falcons, owls). Due aims of increasing and identifying the qual-
to historical pest-resistance problems, resis- ity level of fruit produced in order to
tance management is also an important part increase exports, to have access to the most
of the guidelines. exacting markets, to create a positive image
In spite of the production and marketing of the region (in terms of human-health and
pressures, the introduction and application environmental-protection issues) and to pre-
of IP according to the AGRIOS programme serve the natural resources of the region. The
met many difficulties in South Tyrol, since programme was based on the philosophy
both growers and packing-houses had a and concepts of IOBC/WPRS and its defini-
sceptical and doubtful attitude towards a tions were accepted. The guidelines and the
concept that established precise rules for the field and storage logbooks were established
participants. In 1989, only 929 orchards cov- by a group of more than 50 persons who
ering 1824 ha and 44 packing-houses joined worked for private and public firms or were
the AGRIOS programme, and therefore the independent advisers. The guidelines
IP label was not released. However, by 2001, include the usual sections on grower training
15,337 ha (87% of the total fruit-growing and the options available for the manage-
area) were initially enrolled in the IP pro- ment of orchard environments, sites, root-
gramme and 6954 growers produced stock and cultivar selection, irrigation
732,920 t of integrated fruit (78% of the total scheduling, soil, tree and fruit management,
South Tyrolean pome-fruit production). pest control, postharvest management and
Grower success in the full adoption of the certification requirements.
programme in 2001 was very high, as only Ten growers and 17 companies applied
11% of the total acreage initially enrolled was the programme for the first time in 1995/96
voluntarily dropped by the growers or elimi- on 100 ha of orchards. The area officially
nated after orchard or field-book inspections. under IP increased up to 1200 ha and then
As a result of the application of the pro- has stabilized around 700 ha since 1999/2000.
gramme, fewer residues are present in fruit, However, IP is applied by a great percentage
2 Source: Magdalena, C., Di Masi, S. and Colodner, A., Programa Producción Integrada de Frutas –
Patagonia, http://www.inta.gov.ar/altovalle/prioneupam.html (information on Alto Valle); http://www.
inta.gov.ar/altovalle/ppfi.html (information on IFP); http://members.tripod.com/intecace/ (IFP Directives).
Apples - Ch.21 21/3/03 3:03 pm Page 546
of growers without being certified. The IP and these developed chapters for the IFP
methods used in Patagonia do not show any manual, covering site selection, rootstocks
economic differences over conventional and cultivars, soils and nutrition, water
methods. In fact, growers have not obtained management, understorey management, tree
any economic advantages in the short term. management, spray application, pests, dis-
However, they mention other advantages, eases, orchard environmental quality, indus-
such as the small environmental impact and try operations, cleaner production, grower
the increase in biodiversity. The programme training and auditing.
is not subsidized and it is audited by an The IFP pest- and disease-management
independent body. strategy was largely developed from earlier
IPM research findings and became the
initial focus for IFP implementation. Prior
21.7 Case-study 3: Integrated to the adoption of the IFP programme, the
Production for Apples in New Zealand3 pest-control strategy was based on a regular
schedule of broad-spectrum organophos-
By J.T.S. Walker. HortResearch, Private Bag phate (OP) insecticides, primarily for the
1401, Havelock North, New Zealand. control of tortricid leaf-rollers. The main
goals of the new pest-control strategy were
New Zealand’s 1500 apple growers export to reduce total insecticide use and eliminate
about 300,000 t of apples annually. Over OP insecticides from apple production by
90% of the Class One apples are exported to 2001. To achieve these goals, the pro-
more than 60 countries, mainly to the UK gramme encouraged the use of selective
and Europe (50%) and the USA (25%). The pest-control methods to maximize biologi-
New Zealand industry faces high produc- cal control and the development of
tion and transportation costs compared postharvest pest-removal systems to
with its southern-hemisphere competitors decrease the risk of quarantine-inspection
and has therefore used fruit quality and failures.
new cultivars as points of differentiation to The introduction of IFP in apples had a
maintain orchard profitability. In the mid- dramatic impact on insecticide use nation-
1990s, food safety and environmentally sen- ally. Between 1996 and 2001, the total num-
sitive production systems became ber of applications declined regionally by
important customer requirements in these 42–58%, while OP insecticide use declined
key export markets. However, phytosani- by 90% (azinphos-methyl use declined by
tary issues were also important factors that 97%, chlorpyrifos use by 82% and diazinon
needed consideration before the New use by 90%). Fungicide applications
Zealand fruit industry changed from a pro- declined by 13%. As a consequence, biologi-
duction system that targeted international cal control of some pests (woolly apple-
quarantine standards towards one based on aphid control by Aphelinus mali, European
IFP principles. red-mite control by Typhlodromus pyri and
Implementation of the IFP programme for mealybug control by general predators) has
apples and pears in New Zealand began in become more widely established.
1996. A national IFP Pip-fruit Committee, led As for other IFP management practices,
by ENZAFRUIT, which included technical all growers must have training and certifica-
experts, growers, consultants, consumers tion for the safe handling of agrochemicals,
and the agrochemical industry, developed and spray-application equipment must be
IFP guidelines based on the philosophy and regularly calibrated. The use of residual her-
concepts of the IOBC/WPRS. The committee bicides has decreased markedly and the
also established technical subcommittees weed-free strip in tree lines is now typically
3 Moreinformation: Walker, J.T.S., Manktelow, D.W.L., Wearing, C.H., Lo, P.L. and Suckling, D.M. (2001)
Development of integrated fruit production programmes in the New Zealand horticultural industry.
IOBC/WPRS Bulletin 24(5), 39–44.
Apples - Ch.21 21/3/03 3:03 pm Page 547
below 30% of the orchard area. Responsible 21.8 Case-study 4: Integrated

fertilizer use and irrigation management are Production for Apples in Northern
encouraged and increasing recognition is Oregon (USA)
now given to soil health and the mainte-
nance of soil organic matter in the IFP pro- For the last 10 years federal funding initiatives
gramme. under the Sustainable Agriculture Research
One of the key achievements of the IFP and Education (SARE) grants programme
programme has been its rapid and complete have promoted the adoption of sustainable
adoption by the apple industry. In the agricultural production practices in the USA
1996/97 season, 88 growers joined a pilot (http://wsare.usu.edu/). Similar to IP, the
IFP programme and, by 2000/01, over 90% mission of SARE is to expand the adoption of
of growers participated. IFP is now the min- sustainable agricultural practices that are eco-
imum export standard for New Zealand’s nomically viable, environmentally sound and
apple growers. IFP recommendations are socially acceptable. This considerable invest-
reviewed annually, analysing industry-wide ment in sustainable agriculture has spawned a
pest monitoring and control records, number of producer organizations in the USA
together with fruit-quality data obtained with their own guidelines and eco-labels.
from packing-house grading procedures. However, compared with other fruit-growing
This allowed identification of difficulties regions around the world, especially Europe,
and provided the basis for programme revi- efforts to establish IP programmes for apples
sions to ensure that fruit quality and grow- and other tree fruits with recognized labels
ers’ confidence in the programme was and full certification programmes have so far
retained throughout. During implementa- been limited. Often these programmes involve
tion, growers were also required to attend producers in a limited geographical area and
discussion groups led by trained facilita- are not inclusive of all production practices.
tors. Any issues arising from these groups Their focus has generally been on IPM and
restricting the use of agricultural chemicals,
were collated nationally and resolved by
especially broad-spectrum pesticides.
scientific experts, who then informed all
Examples are the ‘Core Values Northeast’ pro-
facilitators.
gramme (http://www. corevalues.org/) for
Several factors contributed towards the
apples in the north-eastern USA and the
rapid implementation of IFP. One of the most
‘California Clean Growers’ programme
important was having a unified marketing
(http://www.californiaclean.com/) for fruits
structure, where one organization could set
and vegetables.
uniform production standards across the One area that has begun to follow the
whole industry. ENZAFRUIT also assisted European IP model is the Mid-Columbia
implementation by giving a small financial fruit-growing district in northern Oregon.
incentive to IFP growers. But most growers This region produces apples on 610 ha, pears
welcomed the move away from intensive use on 4860 ha and sweet cherries on 2830 ha.
of OP insecticides and found that the IFP Since the early 1990s, efforts have been
programme did not incur significant addi- under way to develop IP programmes for
tional costs, while the fruit quality at harvest apples, pears and sweet cherries in that area.
was comparable with the old OP-based pro- The IOBC guidelines for pome fruits (Cross
gramme. At present, growers enjoy the IFP and Dickler, 1994) served as a template for
programme and they have clear marketing these programmes. Another IFP programme
benefits with customers. The grower-owned in Oregon involves grapes and is known
company New Zealand Pipfruit Limited under the acronym LIVE (Low Input
now manages the programme. An inde- Viticulture and Enology). The grape IFP
pendent agency, on behalf of exporting programme (http://liveinc.org/index.htm)
companies, audits growers’ pesticide use and was developed independently of the other
compliance with the programme, to ensure IFP programmes in Oregon and is the first
the integrity of IFP. IFP programme in the USA endorsed by the
Apples - Ch.21 21/3/03 3:03 pm Page 548
IOBC. The pome-fruit and the cherry IFP been on education and programme develop-
programmes in northern Oregon are at a ment and less on marketing. No mechanism
similar point of development. Initially, the is in place at the present time to certify or
emphasis was on building the infrastructure audit growers. However, fruit growers who
for each programme. This included consti- want to be certified according to production
tuting grower-led IFP committees, which standards similar to the IFP guidelines can
oversee the development, implementation obtain certification through The Food Alliance
and direction of each programme. University (TFA), a non-profit organization dedicated to
research and extension play only a supportive sustainable agricultural practices (http://
role by conducting IFP-related research and www.thefoodalliance.org/). TFA guidelines
grower education. IFP guidelines for pome emphasize three areas: pest management, soil
fruits and cherries were first published in and water conservation and farm labour. In
1994 and they spell out the aims and pre- addition to certification, TFA assists with pro-
ferred practices under IFP (http://www. motion and marketing. It is estimated that
orst.edu/dept/hort/orchardnet/ifp.htm). about 8% of the pome-fruit and cherry
The IFP guidelines are annually updated. acreage in the Mid-Columbia fruit-growing
Pesticides are listed in order of preference district is now certified by the TFA.
and IFP compatibility. In addition, demon- In addition to raising grower know-how
stration orchards throughout the district of fruit production by updating them regu-
serve as vehicles for showcasing and pro- larly about new technologies, the IFP pro-
moting IFP practices such as selective pest grammes in the Mid-Columbia fruit-growing
management without broad-spectrum pesti- district have had a positive impact on horti-
cides; biological control of major fruit pests; cultural and pest-management practices.
pest and disease control based on monitor- This is also evident from the substantial
ing and model forecasts; yield mapping; and decrease of OP insecticide use between 1993
site-specific water and nutrient management. and 1999. For instance, azinphosmethyl use
Educational programmes and materials are on pome fruits has decreased by more than
delivered to growers, consultants and packing- 20% over that time period. With increasing
house representatives through a cooperative use of mating disruption and insect-growth
effort of local grower organizations and the regulators for codling-moth control, even
university. The focus of educational pro- more dramatic reductions in OP use can be
grammes is to support knowledge-based expected in the next few years.
orchard-management decision-making. To
assist growers with irrigation scheduling
and pest and harvest management, weather Acknowledgements
information and pest and disease forecasts
are available via telephone and the Internet. We are grateful to the following colleagues,
This information is updated daily from a who have provided information about the
network of remote weather stations. A com- case-studies: C. Magdalena, S. Di Masi
puterized spray-record system is being put and A. Colodner (Instituto Nacional de
in place for growers to report pesticide use Tecnología Agropecuaria, Argentina), for
on individual blocks. The system will track Case-study 2, and P. Wierer (AGRIOS, Italy)
pesticide use in the district and document and W. Waldner (Südtiroler Beratungsring
the implementation of IFP practices. für Obst und Weinbau), for Case-study 1. We
So far the emphasis of the Oregon IFP also thank J. Walker (HortResearch, New
programmes for pome fruits and cherries has Zealand), who has written Case-study 3.
References
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Apples - Ch.22 21/3/03 3:03 pm Page 551
22 Organic Apple Production – with

Emphasis on European Experiences
Franco Weibel and Andreas Häseli

Research Institute of Organic Agriculture
(Forschungsinstitut für biologischen Landbau (FiBL)), Frick, Switzerland

22.2 Basic Principles of Organic Farming 552
22.3 The Legal Basis 553
22.3.1 Important requirements in organic fruit production 554
22.4 The Extent and Importance of Organic Fruit Production in Europe 554
22.5 Supply Situation and Market Outlook 554
22.5.1 Outlook for the organic fruit market 554
22.5.2 Distribution channels and marketing structures 556
22.5.3 Grading rules 556
22.5.4 Producer prices 557
22.6 Conditions Favouring the Conversion to Organic Apple Production 557
22.7 Site Requirements 557
22.8 Planting Systems 558
22.8.1 Environmental considerations in choosing construction materials 559
22.9 Choice of Rootstock 559
22.10 Areas and Features for Ecological Compensation 559
22.11 Choice of Cultivars 560
22.11.1 Choosing a suitable range of cultivars for commercial organic fruit production
and marketing 560
22.12 Organic Apple Production Requires New and Autonomous Marketing Concepts 562
22.12.1 Grouping apple cultivars into archetypes 563
22.13 Soil Management and In-row Weed Control 563
22.13.1 Sensitivity to weed competition 563
22.13.2 Soil preparation prior to planting 564
22.13.3 Orchard-floor management: controlling ground-cover competition instead of
killing weeds 565
22.13.4 The problem area around the trunk 565
22.13.5 The most interesting new developments on the market 565
22.13.6 The Landurner mechanical hoe 565
22.13.7 Thermal weed-control systems 567
22.13.8 Outlook 568
22.13.9 Mulching the in-row strip 568
22.14 Nutrient Management 568
22.14.1 Important helpers in providing nutrients 569
22.14.2 Fertilizer use 569
22.14.3 Nitrogen fertilization 569
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552 F. Weibel and A. Häseli
22.14.4 Soil-improvement products 570

22.14.5 Foliar feeds and tonics 570
22.15 Thinning 571
22.16 Plant Protection 571
22.16.1 Aims and principles of plant protection in organic fruit production 571
22.16.2 System stabilization as a basic principle 572
22.16.3 Indirect control measures 577
22.17 Organically Approved Disease-control Products 577
22.17.1 Clay-powder preparations 577
22.17.2 Wettable sulphur 577
22.17.3 Copper 578
22.17.4 Coconut soap 578
22.17.5 Lime sulphur 578
22.18 Organically Approved Products for Pest Control 578
22.18.1 Plant extracts, soaps and oils 578
22.18.2 Biological methods 579
22.19 Aspects Of Farm Management in Organic Fruit Production 580
22.19.1 Labour input and direct costs compared with integrated apple production 580
22.19.2 Yields 581
22.1 Introduction the major wholesalers have introduced

organic fruit into their range – e.g. COOP in
Regardless of the production methods Switzerland with their NaturaPlan range,
employed, fruit growing requires a large Bila in Austria and Reve in Germany – has
amount of care. During the long vegetation given the market a strong new impetus.
period, the trees are exposed to many pests The current market share of organic fruit is
and diseases and, since fruit trees are perma- approximately 3–5% of the dessert-fruit mar-
nent crops, any management mistakes can ket in Switzerland, while in Germany it is
continue to have an effect for a number of currently less than 1%. Experts agree that, in
years. Furthermore, the demands on quality the medium term, organic fruit can increase
are very high, since fruit is usually sold with- its market share to 10% (Schmid et al., 1995).
out any processing.
Compared with the conventional pro-
ducer, the organic fruit producer faces extra 22.2 Basic Principles of Organic
challenges and production risks as a result of Farming
the rigorous restrictions imposed upon pro-
duction aids. Over the past years it has been The aims and objectives of organic cultiva-
possible to greatly improve yield stability in tion – applicable worldwide – are as follows:
organic fruit production as a result of the ● to produce high-quality foods that are
substantial progress made in terms of free of artificial and health-damaging
cultivars, production aids and machinery. pesticide residues;
Additionally there have been welcome ● to build and maintain high soil fertility;
developments in the economic environment, ● to maintain natural cycles and processes
owing to lively demand and fair producer that are as closed as is feasible;
prices. ● to promote and conserve biodiversity;
Both the supply of and demand for ● to keep, feed and breed livestock in a
organic fruit have grown rapidly since the manner that observes particular ethical
mid-1990s. Demand continues to outstrip principles;
supply, which shows that consumers increas- ● to work without the use of synthetic aids
ingly value foods that are untreated and free and ingredients, or genetically modified
of pesticide residues. The fact that some of organisms and their derivatives.
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Organic Apple Production 553
(Simon, 1995) still give shape to the organic

standards used today.
The majority of organic fruit producers in
Europe produce to ‘organic’ standards,
rather than ‘biodynamic’ standards. Vogt
(2000) provides an up-to-date historical
overview and a listing of original sources.
Most of the principles and methods
described in this chapter apply to both bio-
dynamic and organic farming. We have
refrained from providing a detailed descrip-
tion of the specific requirements and treat-
ments used in biodynamic fruit production
(biodynamic preparations, consideration of
planetary constellations, composting meth-
ods, etc.) in this chapter, as this would
require a more in-depth discussion of the
underlying philosophy.
Fig. 22.1. Rudolf Steiner (1861–1925), the founder
of biodynamic agriculture.
22.3 The Legal Basis
There are two main organic farming The legal minimum standards for organic
systems: organic farming and biodynamic production, processing and specification
farming, the latter essentially amounting to of organic products are laid down in EU
the former but including additional require- Regulation 2092/91 (EEC, 2000). This regula-
ments and treatments. The foundations for tion also forms the basis for the national
biodynamic farming were laid with a course organic regulations in the individual
of eight highly regarded lectures by the sci- European countries and for the organic
entist, philosopher and founder of the spiri- labels established under private schemes,
tual science of anthroposophy, Rudolf such as Demeter, Bioland, Naturland or BIO-
Steiner (1861–1925) (Fig. 22.1; Steiner, 1924). SUISSE (BIO-SUISSE, 1999). The private
Based on this school of thought, biodynamic inspection and certification bodies are free to
farming, building upon scientific principles, prescribe standards that are stricter than
aims to work with non-physical forces, such those contained in the EU Regulation.
as special compost preparations, which are However, they must not certify to a lower
based on minute doses, or the timing of standard than that set by the EU or by
planting and cultivation on the basis of plan- national legislation. Most European organic
etary constellations (Koepf, 1993). producers are members of a private inspec-
In the German-speaking parts of Europe, tion and certification body, because this has
organic farming (biologisch-organischer clear advantages in terms of marketing and
Landbau) was influenced strongly in the access to information. A detailed comparison
postwar period by Dr Hans Müller and Dr of the most important European sets of
Hans-Peter Rusch. Other pioneers, such as organic standards has been published by
Sir Albert Howard and Lady Eve Balfour Häseli (1996).
(Willer, 1995) in England, as well as Jean To foster global harmonization of stan-
Boucher and Raoul Lemaire in France, dards with the aim of achieving equivalence
enriched the movement with their work for organic products, the International
(Siebeneicher, 1995). Organic farming is Federation of Organic Agricultural Move-
based solely on science, while pursuing the ments (IFOAM), an association of national
same aims as biodynamics. The general organic certifying bodies under private law,
guidelines formulated by Müller and Rusch established its own basic standards in 1980.
Apples - Ch.22 21/3/03 3:03 pm Page 554
The IFOAM Basic Standards (IFOAM, 1998), about 2–4% of national dessert-fruit pro-
which are revised every 2 years, are particu- duction in several of the main European
larly important for countries that do not as fruit-producing countries. While organic fruit
yet have their own organic standards. They must still be termed a niche product, figures
also form the basis from which the IFOAM for relative annual growth are substantial, at
Accreditation Programme operates – a between 10 and 30%. With the market entry
private programme, set up in 1996, which of major wholesalers since the mid-1990s
evaluates and accredits private national cer- and consistently attractive producer prices,
tification programmes. At the public level, the area under organic fruit has increased in
two United Nations (UN) organizations – the many countries. Southern Tyrol (Italy), for
Food and Agriculture Organization (FAO) example, has experienced a major increase.
and the World Health Organization (WHO) – There are numerous biodynamic fruit
adopted standards for plant products growers in Germany, Southern Tyrol and
from organic agriculture in 1999 for the first France, while there are only a small number
time as part of the Codex Alimentarius (an of biodynamic holdings in Austria and
international foods standards programme) Switzerland. Throughout Europe, almost all
(FAO/WHO, 1999). The standards of the of organic fruit production is on holdings
Codex Alimentarius are important for the that have been converted in their entirety,
World Trade Organization (WTO) in assess- with very few exceptions.
ing the equivalence of organic products.
IFOAM holds observer status with the Codex
Alimentarius Commission. 22.5 Supply Situation and Market
The inspection and certification of hold- Outlook
ings is carried out in individual countries at
least once a year by an independent, accred- 22.5.1 Outlook for the organic fruit market
ited inspection company. Certifications for
imports of products into the European Union In an extensive survey among experts in
(EU) are mostly based on the certificates of Switzerland in 1994, survey participants esti-
the accredited certification bodies in the orig- mated the market potential for organic fruit
inating countries, provided these are recog- in 2000 at approximately 10% (Schmid et al.,
nized by the EU. Otherwise certification is 1995), with about one-third of production
carried out by accredited inspection and cer- reaching consumers via wholesalers.
tification bodies within the EU. Today the 10% market share forecast still
appears realistic for the future. Its realization
has been delayed because the development
22.3.1 Important requirements in organic of these markets requires more effort on the
fruit production part of traders and producers than originally
anticipated.
The various sets of standards mentioned The main obstacles are that it is essential
above contain separate sections dealing with for organic fruit to be positioned separately
the rules for organic fruit production. Apart from conventional fruit in retail outlets, and
from the general production standards, the that additional consumer information has to
following prescriptions and recommenda- be provided (see Section 22.12). In order to
tions are of particular importance for organic win over the supermarket customer as a
fruit producers (Table 22.1). buyer of organic fruit, both the inner quality
and the appearance of the fruit have to be at
consistently high levels. Furthermore, the
22.4 The Extent and Importance of price differential to conventional fruit must
Organic Fruit Production in Europe not exceed certain thresholds and it must
be based on transparent economic data
According to a survey carried out by Häseli recorded at every step in the production
(1998), organic fruit production amounts to process.
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Table 22.1. General production requirements for organic fruit growing.
Topic Recommendations Prescriptions
Conversion Most private certification bodies require

the immediate conversion of the entire
farm to organic farming practices
(extension of the conversion period to a
maximum of 5 years is possible). Under
certain circumstances, the EU allows for
partial conversion and organic production
in partial separate units for an unlimited
period
Soil management Preferably permanent ground cover; No herbicides; no clean cultivation

species-rich flora and fauna; competing
vegetation is controlled while
simultaneously protecting the soil and
maintaining a long-lasting natural
ground cover; encourage and maintain
good soil structure and soil biological
activity
Fertilizers and mulch Leave organic material as mulch or No synthetic nitrogen or phosphorus
incorporate by shallow cultivation. fertilizers and no muriate of potash
Prudent use of fertilizers in keeping fertilizers
with requirements, in order to reduce The use of certain bought-in fertilizers
plant protection problems and to (basic slag, crude potassium salts and
improve fruit quality magnesium potassium sulphate), as
well as trace elements and calcium
chloride for bitter pit, require that the
need be recognized by the inspection
authority or inspection body and must
be notified. Fast-acting calcium
fertilizers, such as slaked lime (Ca(OH)2)
or quicklime (CaO), are not permitted
because of their caustic effect on soil
organisms
Production systems, Provide for sufficient light penetration

tree shape and air circulation
Cultivars, Choose cultivars and rootstocks No genetically modified cultivars or

planting stock adapted to location and production rootstocks; planting stock must be
system organically produced (transition period
in the EU until 31 December 2000). A
number of producer associations have
developed relevant guidelines
regulating the details. In 1999 the
Swiss working group of organic tree
nurseries (Arbeitsgemeinschaft
ökologische Baumschulen (AGÖB))
published, together with the certifying
bodies, a detailed compilation of
quality requirements for organic
planting stock. These are as strict as
those for conventional stock (AGÖB,
1999)
Continued
Apples - Ch.22 21/3/03 3:03 pm Page 556
Topic Recommendations Prescriptions
Pruning Prune trees to ‘naturally spreading’

shape and moderate growth; support
physiological balance in trees; remove
any diseased, dead and broken limbs
that harbour pests or diseases
Plant protection Use preventive measures to regulate No use of synthetic plant-protection

pests and diseases, for example: products; use of plant protectants on the
use cultivars that are disease-resistant basis of standardized counts; permitted
and pest-tolerant and that are adapted aids are listed in Annex III of EU
to the local soils and climate Regulation 2092/91; Switzerland has a
Provide habitats for beneficial insects binding limiting list showing the available
and birds, such as strips sown in wild products that have been tested –
flowers and herbs, hedgerows, etc. including the additives – for suitability
with organic standards and their efficacy
under organic conditions
Growth regulation Undertake timely thinning of fruit, Thinning sprays and growth regulators
and fruit thinning either manually or mechanically (e.g. are not permitted
with the Fadengerät – a mechanical
tool for thinning blossoms, described
below)
Quality and grading Strive for the best possible inner These aspects are not covered by EU
quality and food safety regulations and usually the conventional
grading rules are applied. In
Switzerland, more tolerant grading rules
in relation to blemished fruit and fruit
sizes are applied in cooperation with the
wholesale and retail trade
Storage, processing Apply careful processing with No restrictions on atmosphere control in

minimum treatment storage rooms
Some private certification bodies do There are restrictions on processing
not permit juices made from aids. Permitted aids are: agar and pectin
concentrated fruit juice for jams, fresh egg-white and natural
casein for fining, pure-culture yeast,
sulphurous acid for cleaning by
fumigation, gelatin
22.5.2 Distribution channels and marketing 22.5.3 Grading rules

structures
In the EU, generally the same quality stan-
In Southern Tyrol, practically the entire dards apply to organic and conventional
organic fruit harvest is marketed through fruit. In practice, organic apple producers are
wholesalers. The main markets are Germany able to sell class II grades mixed with class I
(70%), central and southern Italy (15%) grades as dessert fruit. Switzerland is devel-
and Great Britain (10%) (Häseli, 1998). In oping its own, nationally applicable rules on
Germany, Austria and Switzerland, there is a sizing for organic fruit. The standards are
similar trend towards marketing through the part of the BIO-SUISSE Standards. They are
wholesale trade (Table 22.2). discussed on an annual basis between organic
Apples - Ch.22 21/3/03 3:03 pm Page 557
Table 22.2. Approximate distribution (%) of organic fruit outlets among different markets (from Häseli,
1998).
Country Wholesalers Retailers Direct marketing
Italy (Southern Tyrol) 95 Of little relevance Of little relevance

Germany 60–65 25 5–10
Austria 70 20 10
Switzerland 60 25 15
fruit producers and traders and revised if be successful both economically and from
necessary. The ultimate aim of these grading the point of view of crop husbandry requires
rules is to guarantee attractive-looking fruit first and foremost highly motivated and
with good flavour and high hygienic quality. highly skilled managers and staff. It is of
Small blemishes are accepted by both traders benefit if the manager’s family is prepared to
and consumers, which ensures that the take some risks, does not shy away from the
intensity of plant-protection measures is kept professional challenge and enjoys living and
at an environmentally acceptable level and is working with a species-rich ecosystem and its
not dictated by ‘cosmetic’ quality demands. many interdependencies. Continuous further
education and training are very important.
It is advantageous if the orchard property
22.5.4 Producer prices has the following characteristics:
Since demand for organic fruit continues to ● Fruit crops, cultivars and production
outstrip supply by a considerable margin and systems are suited to organic production
higher production costs are demonstrable (see and to the marketing channels envisaged,
Section 22.19), organic apples achieve consid- land is available for ecological compensa-
erably higher producer returns than conven- tion and the restructuring of the holding
tional ones. According to a study carried out for organic production is economically
by the Swiss Research Institute of Organic feasible (for example, plant-protection
Agriculture (Forschungsinstitut für biologis- measures and soil management are very
chen Landbau (FiBL)) (Häseli, 1998), organic difficult with high-density plantings).
producer returns in Southern Tyrol, Austria ● Buildings, machinery and workforce are
and Germany are two to three times those for appropriate or else the required adjust-
conventional produce. In Switzerland, the ments can be financed.
average producer returns for organic apples in ● It is possible to restructure marketing
the past 2 years have been twice those for con- channels, where necessary.
ventional ones. Retail prices for organic fruit ● The additional labour requirements in
in supermarket chains are generally between fruit production (for example, for manual
20 and 60% higher than prices for conven- thinning of flowers, mechanical soil culti-
tional fruit. As with the grading rules, the rec- vation, etc.) fit in well with the other farm
ommended retail prices for organic fruit are enterprises including the times of peak
negotiated between producers and traders at labour demand.
the start of a sales campaign and are published.
A pricing scale is set on the basis of cultivar,
quality grade, size class and storage duration. 22.7 Site Requirements
Generally the minimum requirements used

22.6 Conditions Favouring the for site selection for organic fruit production
Conversion to Organic Apple Production are slightly higher than those for conven-
tional production. The limited or, in some
A conversion from conventional or inte- cases, non-existent possibilities for direct
grated to organic fruit production that is to plant-protection measures (e.g. for scab
Apples - Ch.22 21/3/03 3:03 pm Page 558
(Venturia inaequalis (Cke.) Wint.), powdery part of the customer base (for recommenda-
mildew (Podosphaera leucotricha (Ell. & Ev.) tions on production and cultivars, see FiBL
Salm.), sooty blotch (Gloedes pomigena (Schw.) leaflets on commercial organic fruit pro-
Colby), apple-blossom weevil (Anthonomus duction with standard trees (Brunner et al.,
pomorum L.), voles and mice) means that 2000; Häseli et al., 2000)). Unfortunately, the
optimum site conditions are essential, accident statistics show that not enough
including the following: attention is given to accident prevention in
standard-tree orchards.
● Location. Sites in high-rainfall areas with
The bulk of organic dessert fruit is pro-
precipitation of more than c. 1300 mm per
duced on dwarfing trees. The planting densi-
annum are less suitable for organic pro-
ties in modern organic dessert-fruit orchards
duction because of the restricted options
are generally between 2000 and 3000 trees
for plant protection. The proximity of
ha1. High-density plantings with more than
woodlands can lead to additional insect
4000 trees ha1 have been shown to be prob-
problems (apple-blossom weevil, bark
lematic for organic systems in terms of plant
beetles (Xyleborus = Anisandrus dispar F.))
protection and orchard-floor management.
or can impede the drying off of foliage.
Training regimes aim for good air flow both
● Exposure. Sunny, good air flow (to mini-
through the tree interior and between rows,
mize the incidence of fungal diseases),
as well as for unhindered sunlight penetra-
slightly sloping ground (to allow hoeing
tion around fruit on lower limbs and near
or mulching of tree rows), as frost-proof
the trunk. This can be achieved, for example,
as possible.
with a shape that on all sides tapers strongly
● Soil. Must be of at least medium depth;
towards the top. Otherwise fruit on lower
good structure and well-drained (for good
branches and in the centre of the tree are
trafficability); good humus content to pro-
much more prone to disease, particularly
mote high biological activity and suffi-
apple scab and sooty blotch.
ciently quick mobilization of nutrients in
For organic apple production, mostly
the spring, well-balanced soil fertility (to
single-row spindle-bush plantations are
compensate for the limited possibilities
planted. In order to make manual thinning
for foliar feeding or the application of
and the training of the trees easier and also
synthetic fertilizers).
to allow for hail-protection netting, many
organic orchardists limit the height of the
trees to between 2 and 2.5 m. The height at
22.8 Planting Systems which the lowest fruiting branches are
trained is preferably chosen to be slightly
The chosen planting system should aim to higher than in integrated production in order
combine cultivar, rootstock and planting to leave sufficient room for mechanical hoe-
density in a manner that results in ‘moder- ing and for mulch spreaders. Using planting
ately’ growing, productive trees. It is crucial stock with high grafts is of benefit in order
that the system is adapted to the options that the accumulation of soil around the trunk,
available for mechanization and labour which can arise from hoeing or mulching,
deployment on the property. does not lead to self-rooting of the scion
In many organic orchards, dessert apples cultivar. It is recommended that the ideal
are harvested from trees on seedling root- organic planting stock, with a high graft and
stocks. These are mostly old or local culti- a good heading height, is secured by way of
vars, which are used for self-sufficiency and a contract with a tree nursery prior to the
direct marketing. It is well known that the grafting season.
profitability of standard trees (i.e. those on Producers who use the rope machine for
seedling rootstock) for dessert-quality fruit is fruit thinning – a mechanical tool destroying
well below that of trees on dwarfing root- a part of the blossoms by vertically rotating
stocks. However, standard trees are strongly nylon ropes (see Section 22.15) – must try
image-building and are highly valued by a to train preferably short side-branches on
Apples - Ch.22 21/3/03 3:03 pm Page 559
a horizontal plane. Multiple-row plantings habit or too much of a dwarfing effect, each
cannot be managed efficiently because of resulting in unproductive trees. The advan-
problems in orchard-floor management, tages and disadvantages of a number of root-
plant protection and mechanical thinning. If stocks are described in detail in Chapter 5.
a rope machine is to be used, V-shaped Note that, during their establishment years,
orchard systems are also not feasible. the more vigorous rootstocks are not much
more tolerant of competition than, for
example, the M.9 types. Accordingly, the use
22.8.1 Environmental considerations in of a vigorous rootstock in organic production
choosing construction materials cannot compensate for the cost and labour-
intensive control of ground-cover vegetation
The use of modern cropping systems results during the first 3–4 years after planting
in considerable material requirements for (see Section 22.13). When full cropping
stakes, trellises, hail protection and irriga- capacity has been reached, the slightly more
tion. This is also true for organic production. vigorous rootstocks, such as M.26, M.4, M.7
There are a number of possibilities for or MM.106, are more suited to year-round
optimizing the use of such materials from an ground cover in both rows and aisles than
environmental point of view, along with the very dwarfing, such as M.9, and
meeting the requirements of organic produc- extremely dwarfing types.
tion. Instead of using treated wooden posts, For the grower, the plethora of new
trellis from tropical tree species, posts or rootstocks with little-known characteristics
galvanized-metal rods or native hardwoods produced by the many breeding stations is
(such as false locust, oak or sweet chestnut) difficult to come to terms with. Breeders are
could be used. Experience at the FiBL and in paying more attention to disease and pest
commercial orchards have shown that these resistance (e.g. to fire blight (Erwinia
hardwoods can also be used for supporting amylovora (Burr.) Winslow et al.) or woolly
the hail-protection covers, which require a apple aphid (Eriosoma lanigerum Hausm.),
lot of material. A degree of environmental which can only be welcomed by organic
optimization is also possible and desirable producers. However, trials using new root-
through adopting practices that minimize stocks in wholly organic orchards (in respect
the use of metal and plastic components. of plant protection, nutrient management,
However, substantial progress in this regard weed control by hoeing and some competi-
will only be possible when well-anchored, tion from weeds) are still in their very early
drought-resistant rootstocks become avail- stages (Weibel, 2000). Besides improved
able, as these would require neither stakes nutrient-uptake abilities and tolerance of
nor irrigation systems. weed competition, sufficient anchorage with-
out staking is important from the point of
view of the organic fruit grower.
22.9 Choice of Rootstock
In organic fruit production, it is important to 22.10 Areas and Features for Ecological
consider the selection of a rootstock in great Compensation
detail in order to ensure that the rootstock is
well adapted to the site (heavy or light soils, An inherent part of a plan for an organic
shallow or deep soils, wet or dry climate, orchard is the integration of areas for ecolog-
ability to irrigate, replanting on the same ical compensation to maintain and support
ground or virgin ground, etc.). Generally biodiversity. Strips sown in wild flowers and
organic producers use the same or a slightly herbs contribute considerably to encourag-
more vigorous rootstock than is used in con- ing beneficial insects and thus to reducing
ventional orchards. Unsuitable combinations aphid pest species, such as the rosy apple
of rootstock and cultivar can easily result in aphid (Wyss, 1997). In order to control
too vigorous growth, a biennial-bearing aphids effectively, Wyss suggests that 5–10%
Apples - Ch.22 21/3/03 3:03 pm Page 560
of the area be sown in wild flowers and versatile range of cultivars rich in speciali-
herbs. Other features of importance for ties, from summer cultivars through to late
beneficial insects and birds are species-rich keepers. For larger and more specialized
hedgerows, consisting of native species orchards, it is of interest to produce a smaller
(excluding fire blight hosts), ruderal areas range of cultivars that are adapted to the
established with pioneer plants, extensively needs of the wholesale market, thereby sub-
used meadow strips at the fringe of the stantially reducing the workload involved in
orchard, small stacks of branches and heaps planning, orchard care and harvesting.
of stones, nesting blocks for wild bees, A marketing plan that is tailored to the
perches for birds of prey, nesting boxes for a situation in the orchard will make it clear
variety of bird species and so on (Plate 22.1). which cultivars will be suitable and what
The knowledge required to establish and their characteristics will have to be.
maintain hedgerows and wild-flower strips
in an optimum manner is not to be underes-
22.11.1.2 Preferred disease-resistant
timated. For further reading, including prac-
cultivars for organic fruit production
tical management advice, see LBL (1993,
1994a,b,c) and Schmid et al. (2000). All the apple cultivars commercially available
today are more or less susceptible to scab
and require time-consuming, costly plant-
22.11 Choice of Cultivars protection measures, which, nevertheless,
cannot guarantee sufficiently stable yields
The choice of suitable cultivars is of central from susceptible cultivars. The required in-
importance to successful organic fruit pro- tensive scab-control measures are a constant
duction. Because of the limited options for headache for fruit growers. The repeated
plant protection, organic fruit production treatments are problematic environmentally
places particularly high demands on disease (because of soil compaction, energy use and
and pest tolerance or resistance. Addi- stress imposed on beneficial species of fauna)
tionally, management aspects must be con- and in regard to consumer acceptance.
sidered, such as marketing options, available Therefore, except in the most favourable of
sites and the desired farm structure. locations, cultivars that are moderately to
highly susceptible to scab should not be used
in new organic plantings. Table 22.3 shows
the scab resistance of a number of commonly
22.11.1 Choosing a suitable range of
grown apple cultivars. An interim evaluation
cultivars for commercial organic
of the most important cropping and fruit
fruit production and marketing
properties of new resistant cultivars under
organic management based on trials con-
22.11.1.1 Selecting a range of cultivars that
ducted by FiBL, including initial cropping
match marketing opportunities
and marketing experience, is evaluated in
Marketing has a direct influence on the Table 22.4.
choice of cultivars for production. Once a In most of today’s resistant cultivars, scab
marketing strategy has been decided upon, it resistance is based upon the single gene, Vf.
must be adhered to consistently, at least for The weakness of this approach has been
the lifetime of one generation of trees (c. 15 demonstrated clearly by occasional resistance
years), in order to ensure economic success. breakdowns. In coming years, breeders will
Apart from personal preferences and therefore increasingly offer cultivars with
labour availability, it is primarily the location multiple disease resistance. Hence, from the
of the orchard, i.e. the distance to customers outset, an organic orchard should maintain
and traders, that determines whether the options for flexible adaptation of the range of
fruit will be sold to wholesalers or marketed cultivars it produces.
directly to the consumer. On-farm marketing Apart from its degree of pest and disease
can be made much more attractive with a resistance and its taste qualities, a cultivar’s
Apples - Ch.22 21/3/03 3:03 pm Page 561
Table 22.3. Susceptibility of apple cultivars to scab (from Häseli and Bosshard, 1993, and surveys of 17
organic fruit producers 1989 to 1992).
Susceptibility to scab Cultivars
Resistant Various (see Table 22.4)

Low ‘Glockenapfel’, ‘Boskoop’, ‘Jonathan’, ‘Cox’s Orange Pippin’, ‘Delbard Jubilé’
Low to moderate ‘Spartan’, ‘Sauergrauech’, ‘Berlepsch’, ‘Alkmene’
Moderate ‘Kidd’s Orange Red’, ‘Elstar’, ‘Empire’, ‘Gravenstein’, ‘Maigold’, ‘Idared’,
‘Summerred’, ‘Primerouge’, ‘Pinova’a
Moderate to high ‘Jonagold’, ‘Gloster’, ‘Champagner Reinette’, ‘Jakob Lebel’, ‘Delbard Estival’
High ‘Golden Delicious’, ‘Gala’, ‘Meran’, ‘Rubinette’, ‘McIntosh’, ‘Vista Bella’, ‘Goro’,
‘Arlet’, ‘Braeburn’,a ‘Fuji’,a ‘Delblush’a
a Supplemented with more recent experience.
Table 22.4. Summary of 1998/99 interim evaluation of a number of disease-resistant apple cultivars for
commercial organic production on the basis of trials and interlaboratory tests, as well as commercial
grower trials, under organic conditions in Switzerland.a
Archetype (AT) Good impression so far Continue observation Less promising
‘Golden’ AT ‘Resista’ (+ flavour, + ‘Goldstar’ (+ flavour, prone ‘Sir Prize’ (+ good for
appearance, vigorous to bitter pit, large fruit) processing, + yields, + little
growth with poor ‘Delbard Jubilé’ (+ inner tendency towards biennial
branching, powdery quality, large, spreading bearing and not very prone to
mildew, susceptible tree, field resistance to browning of the cut surface of
to rusts) scab only, biennial bearer) the fruit)
Newcomers: ‘Goldrush’,
‘Biogolden’, ‘South Tyrolean
Golden’ types
‘Jonagold’ AT ‘Rubinola’ (+ flavour, ‘Angold’ (+ growth habit, ‘Delorina’ (+ sweetness,

+ appearance, + early + flavour, + Venturia Antonovka + good keeper, + tolerant of
harvest, + self-thinning (VA) resistance, powdery rosy apple aphid, many
(too) well, yield, mildew, large fruit) skin spots, prone to
strong growth) ‘Viktoria’ (+ fruit quality powdery mildew)
(sweet type), + dwarfing tree,
? keeping qualities, ? too
dark)
‘Regine’ (+ fruit quality, + good
keeper, + tolerant of sooty
blotch, poor branching,
yield, tough skin, skin
spots)
Newcomer: CQR-10-T-17
‘Idared’ AT ‘Ariwa’ (+ fruit quality, ‘Florina’ (+ tolerant of rosy ‘Saturn’ (+ appearance,

+ growth habit, + tolerant apple aphid, + yield, + known + growth habit, + frost
of powdery mildew and in the market-place, growth tolerance, keeping qualities,
aphids, fruit thinning vigorous and difficult, flavour, biennial bearer,
required for fruit size powdery mildew, premature fruit fall)
and flavour) moderate eating quality) ‘Priam’, ‘Liberty’, ‘Prima’
(– below average quality)
Continued
Apples - Ch.22 21/3/03 3:03 pm Page 562
Archetype (AT) Good impression so far Continue observation Less promising
‘Rajka’ (+ growth habit, Newcomers: ‘Lotos, ‘Vesna’, ‘Rosana’ (+ overall,

+ yield, +/ flavour, ‘Melodie’, ‘Ahra’, ‘Ahrista’, keeping qualities)
? keeping qualities, FAW 8027 ‘Reanda’ (+ fire blight,
? too dark) flavour)
‘Nabella’ ( strong growth,
susceptible to scab)
‘Cox’ AT ‘Topaz’ (+ flavour and ‘Renora’ (+ resistant to fire

eating quality, + relatively blight, + dwarfing growth,
well known and accepted inner quality not consistent)
in the market-place, Newcomers: ‘Ecolette’,
greasy, bitter pit, ‘Santana’, ‘Gerlinde’
susceptible to aphids)
‘Resi’ (+ growth habit,
+ flavour, + appearance,
small fruit, thinning
required, stores only
up to 3°C)
‘Gravenstein’ AT ‘Retina’ (+ growth habit, ‘Julia’ (+ summer cultivar,

+ appearance, + flavour, + flavour, + vigorous growth)
keeping qualities, ‘Reglindis’ (+ growth habit,
susceptible to aphids) + flavour, bruises very
easily)
‘Hana’ (little experience,
bitter pit)
‘Nela’ (little experience)
‘Baujade’ (+ quality, looks
like ‘Granny Smith’ but doesn’t
taste as good, ripens very
late)
‘Boskoop’ AT ‘Otava’ (+ growth habit, ‘Rewena’ (+ fire blight, + yield, ‘Relinda’ ( strong growth,
+ yield, + multiple poor branching, sooty flavour)
disease resistance, blotch, appearance, flesh ‘Zuzana’ ( strong, difficult
+ appearance, sooty sometimes too soft) growth, poor disease
blotch, ? too acidic for resistance)
an apple similar in
appearance to ‘Golden’ AT)
a+ = strength, = weakness, ? = to be determined (as of 1999).
tendency towards biennial bearing plays an and others. The bearing habits of new resis-
important role in determining economic tant cultivars are not currently known.
success. In order to be able to cope with the
high workload required for thinning in the
limited time around the flowering period, 22.12 Organic Apple Production
the proportion of biennial-bearing cultivars Requires New and Autonomous
in the orchard should to be kept to a mini- Marketing Concepts
mum. ‘Organic cultivars’ with a strong
tendency towards biennial bearing include Despite the undisputed advantages of disease-
‘Boskoop’, ‘Elstar’, ‘Maigold’, ‘Gravenstein’ resistant cultivars for organic apple
Apples - Ch.22 21/3/03 3:03 pm Page 563
production, producers are faced with the prob- be flexible in choosing a suitable range of
lem that both retailers and consumers have cultivars and therefore also to be consistent
limited knowledge of their eating, cooking and in his or her environmental efforts. It further
keeping qualities, not to mention the ecological creates a simple basis for communication
advantages of these cultivars. Additionally, the with wholesalers or retailers. These, in turn,
trend in the apple market towards globaliza- gain greater flexibility in selecting a range of
tion, with a limited number of globally traded cultivars that meet their customers’ needs
cultivars, strongly counteracts efforts to intro- and in communicating effectively the culti-
duce additional cultivars in generally small vars’ characteristics down the distribution
quantities. Currently available disease-resistant chain (intermediaries, consumers).
cultivars are very short-lived, as new cultivars By grouping the range of cultivars grown
with improved eating, cooking and production in an orchard into archetypes, it will be eas-
characteristics are constantly being bred and ier for the producer to see where there is
put on the market. In order to utilize the room for expansion or innovation in plan-
advantages of disease-resistant cultivars, ning the future range. Table 22.5 shows how
despite the difficult conditions prevailing in the new cultivars suited to organic produc-
the market-place, new marketing concepts tion compare in their flavour, i.e. where each
have to be devised for them. of the cultivars susceptible to disease can be
exchanged with robust cultivars.
22.12.1 Grouping apple cultivars into

archetypes 22.13 Soil Management and In-row
Weed Control
The extensive and short-lived range of culti-
vars on offer provides a complexity that both 22.13.1 Sensitivity to weed competition
producers and the market are seeking to
resolve. One way of consolidating and thus Fruit trees in commercial orchards are fairly
simplifying this complexity is to sort the culti- shallow-rooted. Young trees, in particular,
vars into a few, defined groups (Table 22.5): at spread their roots mostly in the topsoil layer,
the retail level the groups are ‘archetypes’, which is rich in nutrients and organic mat-
which are then further simplified as ‘flavour ter. If this zone is also utilized by other
groups’ at the consumer level (Weibel, 1995a; plants, the fruit trees will have to compete
Weibel and Grab, 2000). The definition of the for nutrients and water, which – depending
archetypes is based on the flavour and appear- on the severity of competition – can lead to a
ance of well-known commercially important significant reduction in performance. This
cultivars. For example, the ‘Golden archetype’ results not only in short-term yield reduc-
combines all yellow, large, smooth-skinned, tions but also in less growth on branches
mild- to sweet-tasting cultivars. There are bor- and fruiting spurs, less root mass and a
derline cases in this classification and they reduction in both quantity and quality of
may have to be revised in step with experi- fruit buds, which are so important for the
ence. Grouping into flavour groups then com- following year. Hence, strong weed compe-
bines the archetypes ‘Golden’, ‘Jonagold’ and tition often only has an impact on the yield
‘Idared’ in the flavour group ‘mild to sweet’. in the following year. These complex and
The ‘Cox archetype’ and ‘Gravenstein arche- time-lagged linkages require skilful methods
type’ are described as ‘spicy, slightly acidic’ of weed control and are a major challenge to
and finally the cultivars in the ‘Boskoop arche- fruit growers.
type’ are grouped as ‘predominantly acidic, Soil management in organic fruit produc-
spicy’. The cultivar name is, of course, stated tion has two principal aims: to support and
at each level but the information on flavour maintain soil fertility (biological activity and
becomes predominant. physical stability); and to supply the trees
The grouping of cultivars into archetypes with nutrients and water in appropriate
gives the producer the necessary freedom to quantities and at the appropriate time (basic
Apples - Ch.22 21/3/03 3:03 pm Page 564
Table 22.5. Grouping of apple cultivars into six archetypes and three flavour groups.a
Cultivars suited to
organic production,
listed in Flavour group
approximate Additional (colour and text
Archetype (AT) Definition order of ripeningb cultivars on packing label)
‘Golden’ AT Yellow, big, ‘Resista’, ‘Delbard ‘Golden Delicious’

smooth skin, low Jubilé’, ‘Goldrush’,
in acidity, ‘Goldstar’
predominantly
sweet
‘Jonagold’ AT Like ‘Golden’ AT, ‘Rubinola’, ‘Angold’, ‘Gala’, ‘Arlet’,

but red ‘Viktoria’, ‘Delorina’, ‘Jonagold’, ‘Delblush’,
‘Regine’, ‘Pinova’ ‘Maigold’, ‘Fuji’, YELLOW
‘Pink Lady’, ‘mild to sweet’
‘Braeburn’
‘Idared’ AT Medium-sized to ‘Ariwa’, ‘Rajka’, ‘Saturn’, ‘Fiesta’,

big, smooth skin, ‘Santana’, ‘Idared’, ‘McIntosh’, ‘Spartan’,
mild, taste balanced ‘Reanda’, ‘Florina’ ‘Berner Rosen’,
in sugar and acidity ‘Rosana’, ‘Jonathan’,
‘Empire’, ‘Gloster’
‘Cox’ AT Medium-sized to ‘Alkmene’, ‘Liberty’, ‘Berlepsch’,

small, rustic ‘Discovery’, ‘Cox’s Orange
appearance, ‘Kidds Orange’, Pippin’, ‘Kanada
spicy–aromatic, ‘Resi’, ‘Topaz’, Reinette’, ‘Elstar’,
slightly acidic ‘Renora’ ‘Rubinette’
RED
‘Gravenstein’ AT Early, for fresh ‘Julia’, ‘Retina’, ‘Klarapfel’, ‘Vista ‘spicy,
eating, juicy, ‘Primerouge’, Bella’, ‘Jerseymac’, slightly acidic’
crunchy, slightly ‘Reglindis’ ‘Summerred’,
acidic ‘Gravenstein’,
‘James Grieve’,
‘Delbard Estival’,
‘Granny Smith’
‘Boskoop’ AT Pronounced ‘Boskoop’, ‘Iduna’, GREEN

acidity, also for ‘Rewena’, ‘Otava’ ‘Glockenapfel’ ‘predominantly
cooking and baking acidic, spicy’
a As of 2 September 1999 (Organic Fruit Production Commission of BIO-SUISSE/F. Weibel, FiBL).
b Underlined = scab-resistant.
fertilization, supplementary fertilization, soil-management and soil-restoration mea-

control of weeds/cover crops and irrigation). sures are, therefore, central to any holistic
and efficient soil-management strategy.
One to two years prior to a new planting,
22.13.2 Soil preparation prior to planting the condition of the soil should be assessed
carefully so that corrective measures can be
It is almost impossible to correct major soil taken in time. At this stage, soil improve-
deficiencies in existing orchards since, for ments can be carried out in the most efficient
example, the subsoil cannot be broken up way. Approaches that can be taken include
with a deep chisel-type plough. Preparatory the following:
Apples - Ch.22 21/3/03 3:03 pm Page 565
● Assess the soil in terms of soil horizons, the use of ground covers has almost been too
possibly a spade-sample diagnosis and generous. Many organic orchards suffer from
soil tests. too much competition from ground cover.
● Carry out appropriate soil-improvement
measures (e.g. drainage, chiselling fol-
lowed by immediate stabilization by 22.13.4 The problem area around the trunk
sowing deep-rooting plants, green-manure
cover crops, increasing organic-matter There are a number of different options for
content – for example, by adding well- soil management in organic fruit production
rotted compost, etc.) and, where required, (Table 22.6). The suitability of a method is
add brought-in, organically approved, determined above all by two criteria: (i) how
slow-release fertilizers to achieve a good well the method is adapted to a specific soil;
nutrient balance (Ca, K, Mg, P). As a and (ii) how far the area around the trunk is
guide, the ratio of exchangeable Ca : K covered. Any type of work carried out
should be approximately 10 : 1 and the around the trunk and the stake is extremely
K : Mg ratio 2 : 1. If the K : Mg ratio is precarious, be it mowing or hoeing, manual
higher than 6 : 1, up to 90 kg MgO ha1 work or machines. Working too close to the
can be applied annually (Strebl, 2000). tree often leads to serious injury to trunks
Amounts of more than 1 t quicklime and roots, while working too far away from
(CaO) on light soils and 1.5 t on medium the trunk means that ground-cover plants
to heavy soils should be split and applied can form thick vegetative mats around the
over 2 years. This fertilizer should be tree. This is where voles and field mice find
applied in the autumn during dry ideal cover. Other important considerations
periods. are potential impacts of hoeing passes on soil
● The future soil-management system structure and questions of cost-effectiveness.
should be planned with optimum mecha-
nization in mind and the orchard layout
and planting system adapted accordingly. 22.13.5 The most interesting new
developments on the market
22.13.3 Orchard-floor management: Following a period of approximately 5 years

controlling ground-cover competition of stagnation in the development of methods
instead of killing weeds and machinery for soil management without
herbicides, pioneering new developments
Ground-cover plants fulfil an important once again began to appear on the market
function, as their root activity improves the from 1995 onwards (Plate 22.2). However, in
physical structure of the soil and enhances order to optimize sustainable soil manage-
soil biological activity. Ground cover stabi- ment, it is advisable for orchard management
lizes the orchard ecosystem by providing flexibly to combine a variety of the methods
habitats for additional species of insects, available, based on the individual situation
spiders, etc. Therefore the orchardist should and the ability to cooperate with other fruit
not ask whether it is appropriate to tolerate growers.
ground cover at certain times, but rather
when it might be necessary to temporarily
suppress it. For a long time, orchard soil 22.13.6 The Ladurner mechanical hoe
management was dictated by the ‘gardening
sentiment’ of wanting to control weeds The prototype of this cultivator was built
absolutely and no weed was allowed to dis- by Bruno Brugger, an organic fruit grower
figure the tree row throughout the seasons. in Friedrichshafen, Lake Constance. The
In organic fruit production, where no herbi- Ladurner Company in Southern Tyrol fur-
cides are used and where, until recently, ther developed it for commercial produc-
there were hardly any alternative treatments, tion (Plate 22.2a). The mechanism,
Apples - Ch.22 21/3/03 3:03 pm Page 566
Table 22.6. Different methods of weed control available in organic orchard production.
Method Suitable conditions Advantages Disadvantages
Mechanical hoes Site not too steep; only Quite universal; Can have negative
on soils with stable improves impact on soil structure
structure N mineralization in (allow for regular
springtime regrowth of ground cover
to revitalize soil structure);
area around tree trunk
needs to be hand-hoed
once or twice a year
Ladurner mechanical Where soil is too heavy Efficient and precise Expensive (co-operative
hoe (Plate 22.2a) for other cultivator and even under difficult use)
sward too dense conditions; easy to use
Humus-Planet Requires somewhat Also available as In dense swards, weeds
(Plate 22.2c) easier conditions bilateral attachment can wrap around and
(soil, sward) than and with hydraulic widthplug the machine, on
Ladurner’s hoe adjustment; in that heavy soils a smear
configuration, highly layer may form; ‘wavy’
efficient on light soilsdemarcation of the traffic
way
Eurocomex Jolly Requires somewhat Simple machine that can Slightly less efficient than
easier conditions also open up relatively Ladurner; more difficult
(soil, sward) than dense swards to adjust and to steer
Ladurner’s hoe
Undercutters (Clemens, Only on light soils with Simple machine that Not satisfactory on heavy
Müller and others) a relatively open sward gives good results on soils and in dense
(Plate 22.2b) light soils; high speeds swards
are possible (up to
8 km per hour)
Disc ploughs (e.g. Spedo) More for lighter soils Two operations, which Only moderate results in
(Plate 22.2d) with a good till can be phased: the vicinity of the trunk;
drawing soil away with furrows form in clayey
feeler-support wheel, soils
2–3 weeks later throwing
soil back with fixed
discs and at high driving
speed
Sandwich system Can be used A very simple, Both mechanical hoe

(Plate 22.2g) universally inexpensive cultivator is and establishment and
sufficient; easy to use, management of the
harmless to trees and in-row strip have yet to
soil; combination with be improved for
mulching machines and commercial application
other devices. If aisles
carry vegetation,
additional beneficial
insect habitat is provided
Flame weeders Where cultivators cannot Good impact on Only limited effectiveness
(Plate 22.2e) be used because of emerging weeds; no with perennial weeds;
heavy soils, or used in negative impact on soil energy consumption;
rotation with cultivators structure (ideal open flame can easily
in order to relieve strain supplement to hoeing) damage lower branches
on soils and bark of tree trunk
Apples - Ch.22 21/3/03 3:03 pm Page 567
Method Suitable conditions Advantages Disadvantages
Mulching the in-row strip Cultivator cannot be Balancing effect on soil Requires a lot of material
used because of site water and nutrient and is expensive; some
conditions or for situation; prevents manual labour
operational reasons; erosion and soiling of (weeding/mowing)
mulching material is fruit; can improve remains; control of voles
locally and cheaply physical and biological and mice more difficult
available; soil improve- condition of soil
ment (increase in
organic matter desired)
Mulch sheeting (free- Where mice and voles Always good initial Laying, mulching of
draining, heavy-duty) are not a problem; growth; a good traffic way, cleaning,
(Plate 22.2h) experience in handling long-term solution repairs and vole and
sheeting where it works mice control all cause
problems; hardly any soil
improvement; waste
disposal problem
Bark mulch Only on very free- Improves organic-matter Ligneous content should
(woodchips are not draining soils; where content and soil be low; can be
to be recommended!) addition of organic biological activity; a overgrown by
(Plate 22.2i) matter is required; layer 10 cm thick lasts stoloniferous plants
where there are few 3–5 years; application
problems with equipment for hire
perennial weeds
Rape straw Only where soil is Good weed suppression; Only effective for
free-draining and where encourages earthworms 1.5–2 years
there is a potassium and thus improves soil
deficiency (200 kg structure and biological
K ha1 or more) activity
operational comfort and performance are 22.13.7 Thermal weed-control systems

robust. The results are convincing, even in
situations with dense growth and on heavy A number of different types of thermal
soils (i.e. conditions under which it is diffi- weed-control systems are available. There
cult to hoe but which are quite common in are variants using several torches with an
fruit production). The hoeing is carried out open flame; systems that pass over the
by two rotors with three tines each. The ground with glowing, downward-pointing
front rotor, which is slightly smaller, is con- emitters; and even combinations of open
trolled by a sensor wand and works pre- flames for in-row weeding and infrared
cisely around tree trunks and stakes. All emitters for the remainder of the area (Plate
soil-moving parts are attached to a side- 22.2e). Branches close to the soil line and the
mounted arm moving parallel to the bark of the tree trunk can quite easily be
orchard row. This floating position, with damaged, particularly where flame weeders
lever arms of minimum length and within are being used. Systems that work with hot
view of the driver, allows for optimum steam are still at the experimental stage. By
guidance and control of tillage depth and their nature, thermal weed-control systems
for straight driving. The significant cost of are not as effective in areas of older, dense
this cultivator is offset by the fact that it will and matted weed and grass covers, since the
only have to be used between four and six heat cannot penetrate down to the lower
times a year, so it is perfect for cooperative growing points of the target plants. As long
use. as weeds and grasses are still germinating or
Apples - Ch.22 21/3/03 3:03 pm Page 568
are young, thermal techniques are an effi- 22.14 Nutrient Management

cient alternative to hoeing and are less dam-
aging to soils. Energy consumption has been As in other production systems, the aim of
much reduced in the more modern systems. fertilizer use in organic fruit production is to
It is advisable to combine thermal control provide nutrients at the correct times and in
and mechanical hoeing as, over time, flame- optimum quantities, in order to achieve a
weeding leads to monoculture grass covers. good performance of the trees in terms of
both quantity and quality. It is obvious that
fertilizer use can only be one of many
22.13.8 Outlook measures in establishing trees that are physio-
logically balanced and yield top-quality
The progress made in the development of fruit. For example, if excessive vegetative
weed-control systems is most welcome. growth is promoted, the same yield per area
However, the improved quality of the will be of lesser quality than if it came from
machines brings with it increased costs. FiBL moderately growing trees, because, as a
in Frick, Switzerland, has therefore been result of strong calcium uptake by shoots,
working since 1995 on developing an eco- the fruit eventually store less calcium than
nomic alternative, the ‘sandwich system’ those from trees with moderate growth.
(Schmid and Weibel, 2000; Plate 22.2g). In Only when pruning, thinning and fertilizing
this system, only two strips, approximately programmes are in balance can such prob-
50 cm wide at a distance of about 12–20 cm lems be overcome. However, if the trees are
each side of the tree trunk, are cultivated. starved of nutrients, their assimilation effi-
From trunk to trunk a narrow strip of vari- ciency and vitality declines, resulting in
able width (25–50 cm) with low-growing lowered disease resistance and fruit buds of
vegetation remains uncultivated. The two lesser quality, which in turn lead to lower
cultivated strips are hoed with a very simple yields, biennial bearing and a decline in fruit
and thus inexpensive device without any quality. In organic fruit production, soluble,
separately powered parts. With the ‘sand- fast-release fertilizers cannot be applied and
wich system’ the overall area of ground that foliar feeds are only permitted where the
is kept free of weed competition is as large as need has been demonstrated. A shortage of a
in a clean-cultivation system and in theory particular nutrient, be it as a result of limited
the competition the fruit trees are facing availability in the soil or poor conditions for
should not be notably different. However, uptake (for example, because of soil com-
the ‘sandwich’ method cannot yet be recom- paction or waterlogging), cannot therefore be
mended. Both the mechanical hoe and the corrected in the short term.
establishment and management of the in- In order to provide the trees with suffi-
row strip need to be improved and, even cient nutrients, both in quantity and at the
though both trees and yields have developed correct times, the fertilizing programme
well so far, the results of specific trials are followed by the organic orchardist strives
not yet available. A positive impact on the primarily to support soil fertility in a holistic
in-row soil structure and additional benefi- manner. This means that, apart from provid-
cial insect habitat in the aisles’ species-rich ing sufficient and balanced amounts of nutri-
flora are to be expected but have not yet ents, the soil structure and soil microbial
been demonstrated. activity are enhanced and maintained.
A sufficient release of nutrients from
organic and organically approved mineral
22.13.9 Mulching the in-row strip fertilizers can only be achieved when high
soil microbial activity allows for the mineral-
Mulching can give good results, particularly ization of fixed nutrients, making them avail-
on light soils and those that are dry in sum- able for root uptake. For this reason, organic
mer or are in need of organic matter (Weibel, standards do not allow, for example, the use
1995b; see Table 22.6). of fast-acting lime fertilizers, such as CaO or
Apples - Ch.22 21/3/03 3:03 pm Page 569
slaked lime (Ca(OH)2), with their caustic phosphorus (see overview by Marschner,
effect on the soil fauna. 1995). Phosphorus deficiency practically
The soil microfauna itself requires suffi- never occurs in orchards that have a healthy
cient energy supplies, in the form of organic- soil (Quast, 1986).
carbon sources, as well as a continuity of
pore space in the soil up to the soil surface,
to ensure gas exchange for respiratory activ- 22.14.2 Fertilizer use
ity. The ideal is a deep, friable soil similar to
a forest soil. Soils with ample clay–humus Nutrient-withdrawal figures adapted to the
crumbs (friable structure) are characterized current yield situation provide a good basis
by a large internal surface, providing ample for calculating the required amounts of fertil-
cation- and anion-exchange sites for nutri- izer. Nutrient withdrawal with fruit is rela-
ents and a high water-holding capacity. tively small and, at 30 t of apples ha1,
Balanced moisture availability in turn bene- amounts to 20 kg N, 12 kg P, 45 kg K, 12 kg
fits nutrient turnover and water uptake. In Ca and 4 kg Mg (see Chapter 12). The
such soils the nutrients, their availability, fertilization programme needs to be based
moisture and gas exchange are not just suffi- on soil-analysis results and, where deficiency
cient but also well stabilized (buffered) symptoms occur, possibly on leaf-analysis
through numerous interrelationships. Such a results. In Swiss fruit production, the fertil-
stable environment protects roots from izer recommendations take into account the
stressful conditions and ensures that the physiological (im)balance of the orchard
trees have access to water and nutrients in by factoring in additional aspects (shoot
keeping with their physiological require- growth, cessation of shoot growth, fruit-bud
ments. The need for corrective applications formation, yield, physiological problems,
of single nutrients is thus minimized or elim- rootstock, soil depth, soil composition and
inated. In order to reach this ideal, organic organic-matter content).
farming works primarily with organic (farm-
yard) manures or green manures.
22.14.3 Nitrogen fertilization
22.14.1 Important helpers in providing Supplying nitrogen to apple trees is a prob-

nutrients lem not so much of quantities but of correct
timing. The important phase of nitrogen
Earthworms are indispensable for the demand begins before flowering time (i.e.
creation of macropores and clay–humus during a time when the soil has hardly
complexes. The roots of the apple trees, warmed up and nitrogen supplies from
which tend to concentrate in the topsoil soil-borne nitrogen mineralization are low).
level, can penetrate the more dense and However, during the formation of fruit
less nutrient-rich subsoil levels through the buds and at the rosette stage, the tree can
worms’ humus-clad tunnels. This increases supply approximately half of its nitrogen
the utilizable root space. Of the many requirements from its own reserve proteins
soil (micro)organisms with their variety of (arginine and asparagine in particular)
functions, the asymbiotic and associative (Faby and Naumann, 1986, 1987; Sanchez et
nitrogen-fixing bacteria are of particular al., 1990). A timely supply of the full nitro-
importance. In monocrop leguminous gen requirement can therefore well be
stands, the nodule bacteria can fix 200–400 achieved without actually applying nitro-
kg nitrogen ha1 year1 from the air. gen fertilizers – for example, by hoeing the
Mycorrhizal fungi are also important for in-row winter ground cover before its
feeding trees in organic fruit production. growth commences (roughly in early April).
They are closely associated with the active In this way, the ammonium nitrogen and
root hairs and, in exchange for assimilates, nitrate nitrogen stored in the sward during
they provide the tree predominantly with the winter, as well as the small amount of
Apples - Ch.22 21/3/03 3:03 pm Page 570
freshly mineralized nitrogen, are made fully 22.14.4 Soil-improvement products

available to the trees.
If nitrogen is applied as young composted The topic of nutrient management also cov-
manure (3–4 months old) or as non-caustic ers various soil-improvement products, such
slurry, these applications should be made as different types of rock dust (e.g. silica-rich
early in the year (February–March in the volcanic rock dust), seaweed products, prod-
northern hemisphere) because of the slow ucts based on brown coal and even so-called
nitrogen mineralization. The maximum ‘energy-informed’ quartz sand, etc. Unfor-
application must be limited by the potassium tunately, the soil-improving characteristics
demand of the orchard, as otherwise physio- of these products have rarely been demon-
logical damage could be induced by an strated by independent research, even
oversupply of potassium. An application of though there are regular reports from com-
30–40 m3 or 20–30 t of composted manure mercial growers that they are happy with
ha1 contains about 50 kg N but also 140 kg K. them. In the case of seaweed meals, for
Organic standards also permit the use of example, this is quite conceivable, as they
certain fast-release liquid and solid nitrogen contain calcium, organic compounds and
fertilizers. These include horn meal and horn many micronutrients in a readily available
shavings (12–14% N; horn meal takes effect form, which can spur soil microorganisms as
within 10–14 days, while horn shavings take well as plant roots into action.
8–10 weeks), castor-oil meal (5–6% N), blood
meal (12% N), vinasse (3.5% N, a by-product
of sugar-beet processing), hair or feather 22.14.5 Foliar feeds and tonics
meal (13% N), bone meal (6% degreased),
Organic production strives to feed plants in a
amino acid solutions (55% amino acids and
harmonious manner by way of a healthy soil
peptides, 9% organic N) and oil-seed pulp
and a healthy root system. Foliar feeding
(4.5–8.5% N). These substances can be partic-
is akin to fighting symptoms and should
ularly useful to fill the springtime nitrogen
only be used as an emergency measure.
deficiency mentioned above. However, trial
Consequently, foliar feeds may only be used if
results also suggest caution. Using these
a deficiency has been notified to the certifying
fertilizers can easily induce nitrogen im-
body; their use must be registered. The same
balances and can give rise to a decline in
conditions apply to the use of calcium chlo-
quality. Soil fertility that is less than optimal
ride for the prevention of bitter pit. The fol-
cannot be and should not be compensated for lowing products may be used: magnesium
by means of fast-release nitrogen fertilizers. sulphate (Epsom salt) and preparations con-
Supplementary phosphorus and potassium sisting of sulphates and chelates of iron,
fertilizers should only be applied if a need has boron, manganese, zinc and molybdenum.
been ascertained through soil testing. The fol- There are not as yet sufficient trial results to
lowing phosphorus fertilizers are permitted assess the efficiency of foliar feeding with
under organic management: soft ground-rock amino acid or vinasse products (see above)
phosphate, basic slag and aluminium calcium for the short-term alleviation of nitrogen defi-
phosphate. Permitted potassium fertilizers ciencies or for a general improvement of fruit-
include non-chlorinated potassium salt (e.g. bud quality. The same is true for so-called
kainit), sylvinite, potassium sulphate and tonics. Our recommendation is: (i) in the case
potash magnesia (sulphate of potash magne- of symptoms indicating nutrient deficiency, to
sia, 28% K2O, 9% MgO). immediately carry out a leaf analysis and to
Permitted liming products include calcium send in both affected leaves and leaves of nor-
carbonate of natural origin (e.g. chalk, marl, mal appearance of the same cultivar (as a
ground limestone, calcareous marine algae), control); and (ii) optimize soil-nutrient sup-
phosphate chalk (gypsum), sugar-factory ply in accordance with soil-analysis findings
lime, magnesium and calcium carbonate (e.g. and apply foliar feeds in accordance with leaf-
magnesia chalk, ground magnesium lime- analysis findings. Some trees should be left
stone). untreated as controls.
Apples - Ch.22 21/3/03 3:03 pm Page 571
22.15 Thinning and the tractor speed. Commercial orchards

are already fine-tuning this method, based
The aims of thinning in organic fruit produc- on experience, with a pass on just one side or
tion are exactly the same as in integrated or with very gentle but repeated passes.
conventional production (see Chapter 16). This mechanical-thinning tool primarily
Thinning measures aim to reduce the num- catches blossom clusters on the periphery of
ber of fruit per tree to such an extent that all the tree canopy, which are favourable for
remaining fruit are of optimum quality and fruit development, while those with a ten-
that, at the same time, good fruit develop- dency towards producing poor quality in the
ment is encouraged and biennial bearing is centre of the tree tend to escape. Therefore,
thus avoided. As in conventional production, because of its design, the device is only suit-
an early thinning date is desirable – mouse- able for very slender tree types with good
ear stage until blossom – because of greater light penetration into their centres (slender
effectiveness in preventing biennial bearing. spindles). Apart from the flowers, the nylon
Blossom thinning cannot substitute for fruit strings can also destroy rosette leaves, spurs
thinning after June drop, which is important and the bark of branches. These side-effects,
for quality-fruit formation (Bertschinger which are also unfavourable from the point
et al., 1998). Understandably, organic of view of assimilation capacity and disease
orchardists take a very careful approach to transmission, have to be minimized by way
selectively eliminating flowers, as not only of an observant use of the machine (e.g. use
frosts but also pests, such as apple-blossom before the unfolding of the rosette leaves)
weevil (A. pomorum L.), apple sawfly and a tree-training regime adapted to the use
(Hoplocampa testudinea Klug), winter moth of the machine (few vertical branches). The
(Omophtera brumata L.), the moth Grapholita Swiss Federal Research Institute for Fruit
lobarzewski Nowicki and diseases such as Production, Viticulture and Horticulture
brown rot (Monilia laxa (Ehremb.) Sacc. and (Eidgenössische Forschungsanstalt für Obst-,
Monilia fructigena Pers.), blossom blight Wein- und Gartenbau) has produced an
(Pseudomonas syringae van Hall.) and early- information leaflet on the use of the
season scab (V. inaequalis (Cke.) Wint.), pose Fadengerät tool (Bertschinger and Stadler,
a much greater threat and take more of a toll 1998).
on blossom and young fruit than would be Manual thinning often takes between 100
the case in conventional production. and, in some cases, as much as 200 man-
At present, organic production standards hours ha1 and therefore not only is incredi-
do not allow the use of any growth regula- bly labour-intensive but also has to be
tors, including applications for thinning. carried out within a very short period. For
Manual and mechanical thinning methods larger fruit producers, manual thinning
are permitted (Plate 22.3). The ‘mechanical without the Fadengerät tool is hardly possi-
method’ has been in use since the mid-1990s ble, for both financial and logistic reasons.
when a resourceful orchardist developed There are a number of different
the Fadengerät (H. Gessler, Friedrichshafen- approaches to manual thinning (Table 22.7;
Hirschblatt, Germany). This mechanical Bertschinger et al., 1998).
blossom-thinning device consists of a tractor-
mounted, vertically rotating axle carrying a
variable number of nylon strings of c. 50 cm 22.16 Plant Protection
length; the angle of the axle is also adjust-
able. As the tractor moves along the tree 22.16.1 Aims and principles of plant
rows between the mouse-ear stage and pink- protection in organic fruit production
bud stage or even later, the rotating strings
damage a proportion of the flowers or cut It is the organic ideal that plant-protection
them off completely. The thinning intensity problems are reduced to a minimum once
is thus determined by the number of strings soil fertility is high and the plant and animal
used, as well as by the speed of their rotation communities are diverse and in harmonious
Apples - Ch.22 21/3/03 3:03 pm Page 572
Table 22.7. Summary of different approaches that can be taken to achieve blossom and fruit thinning of
apples under organic production systems.
Method Suitability criteria, advantages Disadvantages
Relieve tree of blossom on one Suitability depends on cultivar (it Very labour-intensive and work
side only, leave the other side is, for example, more successful has to be carried out within a very
and apply corrective fruit thinning with ‘Boskoop’ than with ‘Elstar’). short time
between June drop and preharvest In ideal cases, one-off one-sided With short-stalked cultivars, for
(Bertschinger et al., 1998) thinning has an effect for a example, fruit clusters that are
number of years too dense can remain on the
It is possible to work with entirely bearing side (crowding, sooty
untrained staff blotch and sawfly attacks, soiling
from earwig nests). Young trees
can produce too much vegetative
growth on the non-bearing side
Thinning to about one blossom Young trees do not become Usually requires specialist staff.
cluster every 25 cm one-sided. Few problems with In most cases thinning is not
short-stalked cultivars carried out thoroughly enough. It
is indispensable to calculate the
required number of ripening fruit
per tree and to count out a
sample of blossom clusters
(Österreicher et al., 1998)
Thinning by stripping buds with Simple to carry out; can be done Relatively little quantitative
the back of the secateurs on the during winter pruning and can be control. Early determination of
topside and underside of the carried out by untrained staff flower set (Floriprog, after Dolega
branches Desirable selection for favourably et al., 1997) is recommended
located buds or spurs
Appropriate winter pruning Good time in terms of seasonal As yet little experience with this
workload method. Quantitative
implementation requires
expertise. Requires early
determination of flower set
(Floriprog)
balance. In reality, and particularly where it tion, sufficiently effective solutions are not
comes to speciality crops, such completely available for every problem that may occur.
self-stabilizing systems are rarely found. It is, therefore, even more important to uti-
Even in ‘well-tuned’ decades-old organic lize the entire arsenal of strategies to pro-
orchards, certain pests or diseases can at mote species-rich biocoenoses, which in turn
times get out of control. Therefore, active stabilize the orchard as a whole system. In a
plant protection is an integral part of organic system that is as self-stabilizing as possible,
farming, including both indirect measures individual pests and diseases have a lesser
that stabilize systems and direct control chance of achieving dominance and of caus-
measures (Table 22.8). ing substantial damage. While this ecological
principle is well recognized in theory, it is
very difficult to realize in delicate crops,
22.16.2 System stabilization as a such as a number of fruit species. In some
basic principle cases this holistic approach can fail because
just one of the factors is not optimized (e.g. if
Plant protection in organic fruit production a cultivar is used that is highly susceptible to
is demanding. Unlike in integrated produc- scab or aphids).
Apples - Ch.22 21/3/03 3:03 pm Page 573
Table 22.8. Important pests and diseases in organic apple production, their significance, options for
control, and discussion.
Problem Significance Control options Discussion
Scab High Indirect. Only sunny, well-ventilated Resistant cultivars strongly on the
sites, well-spaced plants and rise. Susceptible cultivars and use
naturally spreading tree shape; of copper sprays should be done
robust cultivars; aid the away with as quickly as possible
decomposition of spores by (replaced with clay-powder
encouraging soil biological activity, products). Modern forecasting
spreading well-rotted compost in methods aim to limit the number of
autumn, flail-mowing fallen leaves. treatments required (lesser impact
Aim for moderate shoot growth on soils and beneficials)
and early cessation of shoot
growth
Direct. Spray cover must be in Scab has been observed on a

place prior to infection period number of ‘disease-resistant’
(only protective effect) and must cultivars. In order to suppress the
be renewed after heavy rainfall spread of the disease, orchards
> 20 mm; if necessary into wet with disease-resistant cultivars
foliage during prolonged wet should also be sprayed during the
periods. At temperatures < 12°C, primary time for the release of
clay-powder products or copper ascopores (no copper should be
(not immediately before or during used). This can generally be
blossom because of phytotoxicity), combined with the control of
at > 12°C wettable sulphur powdery mildew
Powdery Medium Indirect. Tolerant cultivars (but Many of the cultivars that are
mildew resistance to scab should be a resistant to scab are strongly
priority in the choice of cultivars); susceptible to powdery mildew; in
aid decomposition of spores, such cases, the control of powdery
ventilation and cessation of mildew is particularly important for
shoot growth young trees
Direct. Manual removal of

diseased buds and shoots during
winter pruning and as early as
possible during the growing
season; two to five post-blossom
treatments with clay powder or
wettable sulphur (scab-control
treatments also affect powdery
mildew)
Sooty blotch Medium Indirect. Site selection: planting Significance of sooty blotch is
most susceptible (late-ripening) increasing in organic production.
cultivars into the best-ventilated Major losses often result from
spots. Sufficient air circulation insufficiently targeted application
into the centre of the tree; strategies and unsatisfactory
avoiding low-hanging branches, application methods (sufficient
maintain low ground cover cover with active agent with ample
water and high pressure into the
Direct. Depending on risk of centre of trees is crucial)
infection, varying number of
treatments with coconut soap
(on-site experience, cultivar and
Continued
Apples - Ch.22 21/3/03 3:03 pm Page 574
Sooty blotch— weather) from June onwards;

Continued most important applications in
late summer until 3 weeks
before harvest
Postharvest Low Indirect. Meticulous removal of Despite the lack of postharvest

disorders ‘mummies’ and rotted fruit from treatments, there are usually no
the tree. Avoiding late-season major postharvest problems with
scab infections, fruit damage organic fruit if storage conditions
from pests and contamination are good
through splashing soil or in
packaging. Sticking to optimum Research is being carried out on
picking time. Meticulous the use of microbial antagonists
screening of harvested fruit and
good storage conditions. No
wet fruit into storage
Direct. Direct measures are not

permitted and are unnecessary
Rosy apple High Indirect. Support for beneficials Neem is highly effective and
aphid and rosy and cultivar selection. Avoiding specific in its impact (hardly any
leaf-curling root suckers impacts on beneficials are known)
aphid
Direct. Neem preparations before Even very slight spray drift of neem
bud break. Other sprays, such preparations is highly phytotoxic to
as pyrethrums, quassia or soaps, pears (susceptibility varies among
have less effect and only before cultivars)
leaves are curled (pure contact
insecticides)
Woolly apple High Indirect. Supporting beneficials: Cyclical variations in emergence

aphid (on the overwinter branches with woolly are dependent on the development
increase) aphids that have been parasitized of the woolly apple aphid parasitoid
upon by the woolly apple aphid Aphelinus mali (determined by
parasitoid Aphelinus mali Hald in spring weather conditions)
a sheltered place and release
during warm period in springtime.
Choice of cultivars and rootstocks.
Avoiding overvigorous trees and
damage to trees
Direct. At present no organically

acceptable treatments can be
recommended. Combined
approach of sticky bands at base
of tree to stop movement of
larvae, removing colonies by
high-water-pressure sprays
(possibly with addition of soap
as wetting agent) or brush,
delayed dormant applications of
oil emulsions can be useful
(plan with adviser)
Apples - Ch.22 21/3/03 3:03 pm Page 575
Green apple Low Indirect. Supporting beneficials, Often held in check by beneficials;
aphid (high in organic avoiding overvigorous shoot set high control threshold in order
tree nurseries) growth to minimize impact on beneficials
(exceptions are young trees and
Direct. Pyrethrum, rotenone, recently grafted trees)
soap preparations. Restricted to
spot treatments where possible,
to minimize impact on beneficials
Red spider Low Indirect. Supporting beneficials, Outbreaks particularly in years with
mite (occasionally cultivar selection, avoiding warm, dry weather from May to
(Panonychus high) overvigorous shoot growth, July
ulmi Koch), minimizing impact of spray
two-spotted applications on predatory mites The use of sulphur impedes the
spider mite (minimum amount of sulphur), development of both pest and
(Tetranicus releasing shoots or felt bands predatory mites
urticae Koch), hosting predatory mites in late
apple-rust summer and autumn Orchards with high species
mite (Aculus diversity and physiologically
schlechtendali Direct. Most important time to balanced trees do not exhibit this
Nal.) control red spider mite is before problem
oviposition, just post-blossom.
Soap preparations with large
amounts of water, possibly
repeating sprays based on counts
Apple- Low to high Indirect. Avoiding locations in Rarely a main cause of reduced
blossom (in the vicinity the vicinity of forests and yields. Can be a useful ‘thinning
weevil of forests, in woodlands agent’ in years when apple
years with little blossom is abundant
blossom) Direct. Where infestation was
higher than 10–15% in the
previous year and depending
on the result of branch-tapping
sampling during emergence, a
pyrethrum application at > 15°C
can be partially effective
Apple sawfly Medium Indirect. Supporting beneficials,

parasitization has a strong
influence on numbers. Manual
thinning of first affected fruit
prevents attack on further fruit
(larva attacks up to five apples)
Direct. Quassia extract at petal

fall on the basis of the degree of
infestation in previous year; white
sticky traps during blossom and
until fruit set
Continued
Apples - Ch.22 21/3/03 3:03 pm Page 576
Codling moth High Indirect. Supporting bird Granulosis virus is very host-specific
populations, parasitization and thus not harmful to beneficials.
influences appearance Because of its low UV stability and
the moth’s long emergence period,
Direct. Granulosis virus and/or granulosis virus has to be applied
confusion strategies with four or five times where one
pheromones in larger orchards generation emerges and up to nine
times where a second generation
also needs to be targeted. Con-
fusion with pheromones only works
in larger and isolated orchards
Tortrix moth Medium Indirect. Supporting bird populations
Direct. Treatments based on the

degree of infestation in previous
year (monitoring at harvest) with
granulosis virus at the balloon
stage (stage D/E) and early
flowering stage (stage E2)
Winter moth, Low Indirect. Supporting bird Where neem is used to control
tobacco populations, sticky bands to aphids, its partial effect in
budworm control winter moths in controlling winter moths and
mid-October tobacco budworm should be
considered
Direct. Bacillus thuringiensis (Bt)
to control young moth larvae
between pre- and post-blossom
during warm weather > 15°C
(otherwise not sufficient feeding
activity and Bt ingestion)
Laspeyresia Low to medium Indirect. Supporting bird

lobarzewski populations
Direct. Confusion strategy with

pheromones in larger orchards
Voles and mice High Indirect. Large-scale Traps are sufficient for moderate
improvements to habitats for birds vole populations. Satisfactory
of prey, weasels, foxes and cats. control of field mice cannot be
Keeping in-row vegetation and achieved at present with carbon
cover along fences low and dioxide fumigation or trapping
monitoring regularly. Protective
barrier against mice dug into the
ground along the fence line or
around individual trees
Direct. Poisoned baits are not

permitted at present. Traps,
carbon dioxide fumigation using
gas bottles or special equipment.
Sanitation is particularly important
as part of winter preparations
and in the spring
Apples - Ch.22 21/3/03 3:03 pm Page 577
An important criterion in choosing the type ● provision of habitats for beneficial organ-
and intensity of plant-protection measures is isms e.g. with hedges and strips sown in
therefore the question of their short-term wild flowers and herbs (this chapter);
and long-term impact on the self-regulating ● supporting and maintaining soil fertility;
forces within the orchard. ● restrained and balanced fertilization
The differences between the permitted (Chapter 12);
plant-protection options in organic produc- ● planting and training system resulting in
tion and those in integrated/conventional open-structured trees that dry off quickly
production are as follows: (Chapter 15).
● There are some pests and diseases for
which sufficiently effective treatments do 22.17 Organically Approved Disease-
not (yet) exist. control Products
● Only active ingredients and additives to
formulations made from plants and min- 22.17.1 Clay-powder preparations
erals may be used (synthetic substances
are not permitted). Assumed effect: stimulation of induced resis-
● The substances are – with a few excep- tance via phenol metabolism; additionally,
tions – less effective. aluminium ions released in an acidic envi-
● The substances only act on contact (i.e. ronment (pH 3.0–3.5) have a direct impact on
they are not transported in the sap and germinating spores. Clay powder should not
therefore are not systemic). be mixed with alkaline substances, such as,
● The period of effectiveness of the organic for example, seaweed products, granulosis
treatments is mostly shorter. virus or copper.
Effect on scab and powdery mildew:
In effect, these differences are of decisive
extensive trials with clay-powder prepara-
agronomic importance: in converting to
tions have shown effects just as good as
organic production, yield stability cannot be
those of the standard fungicide treatments
guaranteed solely by switching from inte-
with 1.5 kg copper ha1 year1 and sulphur.
grated-production to organic plant-protection
Since clay-powder preparations are more
products. It is indispensable that the reduced
effective than wettable sulphur at tempera-
options for plant protection are supported by
tures under 12°C, they are an adequate sub-
indirect measures, such as cultivar selection
stitute for copper preparations used at these
and the provision of habitats for beneficial
cooler temperatures. Cultivars that are sensi-
organisms.
tive to sulphur and susceptible to rusts are
Adapted application methods and regu-
more tolerant of these preparations than of
larly maintained and well-adjusted tools are
treatments with copper/wettable sulphur.
a prerequisite to success in plant protection.
Trials have also shown a partial effectiveness
For example, spray equipment must be well
against fire blight. Clay-powder applications
maintained, as clay powder, where used,
are not recommended in the summer
increases wear on pumps and nozzles and
because of staining and incompatibility with
can leave enhanced filter residues. The fact
granulosis virus preparations.
that the substances used primarily rely on
In Europe, the experience with ‘film-
contact activity means that complete wetting
coating’ by specially modified kaolin powder
is particularly important.
(registered in the USA as Surround for pest
and disease control) is still insufficient for
22.16.3 Indirect control measures registration and practical recommendations.
The most important indirect measures for

22.17.2 Wettable sulphur
pest and disease control are:
● site selection (Chapter 11); Effect on scab and powdery mildew: the
● selection of robust cultivars (Chapter 4); period of effectiveness is dependent on
Apples - Ch.22 21/3/03 3:03 pm Page 578
temperature and is between 6 and 12 days long. Fr.) v. Arx). For sufficient results, the inten-
At low temperatures not enough control is sity of treatment must be adapted to the risk
achieved and at high temperatures over 25°C of infestation in an optimum manner and
the period of effectiveness is short (only 4–5 proper application techniques are indispens-
days at 30°C) as a result of elevated evapora- able. Coconut soap cannot be mixed with
tion rates. With rising temperatures, application granulosis virus.
rates need to be increased. Depending on culti-
vars and the development stage, phytotoxic
22.17.5 Lime sulphur
reactions can occur, such as sunburn, russeting
and scorching. The fruit are most sensitive from
Lime sulphur is permitted under EU
blossom to T stage (when the calyx cavity and
Regulation 2092/91 for organic fruit grow-
stalk are at right angles). ‘Braeburn’, ‘Cox’s
ing; however, it is not (yet) registered as a
Orange Pippin’, ‘Berlepsch’ and ‘Granny
plant-protection agent in many European
Smith’ are particularly sensitive and the appli-
countries. Trials have shown effects of lime
cation rate must be reduced by 30%. Sulphur
sulphur against scab, powdery mildew and
has a partial acaricidal effect. High application
sooty blotch and for blossom thinning. Lime
rates and short treatment intervals reduce both
sulphur strongly increases the pH on the leaf
pest and predatory mite species.
surface and has a certain curative effect
(Trapman and Drechsler-Elias, 2000).
22.17.3 Copper
Controls scab and partially controls 22.18 Organically Approved Products

European canker (Nectria galligena Bres.) and for Pest Control
bark canker (Gloesporium perennans Zeller &
Childs and Gloesporium album Osterw.). Its 22.18.1 Plant extracts, soaps and oils
biocidal effect results from the copper ions
22.18.1.1 Pyrethrum and rotenone
blocking specific enzyme systems in the
microorganisms’ metabolism. Copper is Pyrethrum is a plant extract from a
more effective than wettable sulphur at Chrysanthemum species; rotenone is extracted
lower temperatures. Admixtures of copper to from the roots of Derris spp. Both are used in
pre-bloom applications and possibly to first apple production to control green apple aphid
post-bloom applications are therefore appro- (Aphis pomi de Geer), pear psylla (Psylla piri L.)
priate. Copper applications during bloom and, where required, apple-blossom weevil
can cause russeting. Copper cannot be mixed (A. pomorum L.). As pyrethrum and rotenone
with clay powder. Copper is a toxic heavy rely on contact activity, pests must be well
metal for the soil fauna and accumulates in wetted with the substances and optimum
the soil. For this reason, many organic regu- application techniques, with a lot of water and
lations and standards specify maximum high pressure, are crucial. Both substances
application rates (e.g. 1.5 kg ha1 per annum have a broad spectrum of efficiency and,
in Switzerland). EU Regulation 2092/91 depending on their time of use, they can dam-
makes provisions for a ban on copper prepa- age certain beneficials. The active agents break
rations in the near future. Copper is avail- down very quickly (half-life of 1–2 days).
able in a number of different formulations
(oxysulphate, hydroxide, chloride and oth-
ers). However, trials have not shown any 22.18.1.2 Quassia
verifiable differences between the effects or
Quassia is a plant extract from ‘bitter wood’
side-effects of these.
(a tropical tree species). It is used in apple
production to control sawfly (H. testudinea
22.17.4 Coconut soap Klug) attacks and is partially effective
against aphids. It acts as a stomach poison,
Controls sooty blotch (G. pomigena Schw.) as well as on contact; the impact on bene-
and fly-speck (Schizothyrium pomi (Mont. ex ficials is low.
Apples - Ch.22 21/3/03 3:03 pm Page 579
22.18.1.3 Neem extract Optimum wetting has to be achieved with

the applications. The impact on beneficials is
An extract derived from the seeds of the
low but repeated applications during the
neem tree, which is widespread, for exam-
summer can cause russeting. Applications
ple, in India. The principal active agent is
during blossom can have a rather unpre-
azadirachtin. The proprietary preparation
dictable thinning effect.
NeemAzal T/S is highly effective against
rosy apple aphids (Dysaphis plantaginea
Pass.) and rosy leaf-curling aphids (differ-
22.18.2 Biological methods
ent species), albeit with delayed action. It
does not control other aphid species. It is 22.18.2.1 Granulosis virus
partially effective against winter moth (O.
brumata L.), tortrix moth (Adoxophyes orana The granulosis virus occurs naturally, is very
F.v.R. syn. Capua reticulana Hb.) and tar- host-specific and as a pathogen does not
nished plant bugs (different Lygus spp.). therefore harm beneficial insects. Granulosis
The impact on beneficials is low. The active virus preparations are used for a highly
agent is absorbed by the leaves and dis- effective control of codling moth (Cydia
persed by way of translaminar flow (the pomonella L. syn. Laspeyresia pomonella L.) and
active agent is partly absorbed and distrib- tortrix moth (A. orana F.v.R. syn. C. reticulana
uted within the leaves). Thus neem is effec- Hb.) on the basis of regionally prepared
tive even if direct contact with the aphids is recommendations for spray schedules.
insufficient (e.g. in the case of curled Granulosis virus is a stomach poison.
leaves). Apart from killing aphids, neem Adding sugar and pine resin extract
also reduces their fertility. With many pear increases absorption and ultraviolet (UV)
cultivars, even very slight spray drift can stability. Since the larvae are not killed
cause severe burns to fruit and foliage. immediately, they can cause small but unob-
trusive and usually well-healed bite marks,
which do not render the fruit unsuitable for
22.18.1.4 Oil sprays dessert-fruit sales. Granulosis virus cannot
Both mineral (paraffin)- and plant-oil prepa- be mixed with coconut-soap or clay-powder
rations are used. Mineral oils are slightly preparations.
more effective but their degradation takes
longer than that of plant oils. Oil sprays are 22.18.2.2 Bacillus thuringiensis
mostly applied at bud break to control scale var. kurstaki (Bt)
insects and are also partially effective
against red spider mite (Panonychus ulmi Used to control winter moth (O. brumata L.)
Koch). However, the latter are more effi- and small ermine moth (Yponomeuta padella
ciently controlled with soap preparations L. and Yponomeuta malinella Zeller); partially
post-blossom. effective against tobacco budworm (Orthosia
species). Bt is a stomach poison, which is
host-specific and therefore not harmful to
22.18.1.5 Soap-based products beneficial insects and predatory mites (e.g.
Soap-based products are manufactured Typhlodromus pyri Scheuten). At temperatures
through saponification of natural fatty acids below 15°C, feed intake is not sufficient.
with alkali lye or a sodium hydroxide
solution (and therefore fulfil the criterion
22.18.2.3 Traps, pheromones and releases
of being natural products). They are mostly
of beneficials
used to control red spider mite and, with
lesser effectiveness, aphids. In order to be Scented or coloured traps are used to moni-
effective against red spider mite, timing is tor the emergence or presence of certain
critical, with applications being made just pests, such as sawfly (H. testudinea Klug) and
post-blossom but before oviposition. different moth species, and to control indi-
Apples - Ch.22 21/3/03 3:03 pm Page 580
vidual species, such as bark beetles (X. dispar 22.19 Aspects of Farm Management in
syn. A. dispar F.), clearwings (Synanthedon Organic Fruit Production
myopaeformis syn. Thamnosphecia myopae-
formis Borkh.) and European goat moth At present, few detailed farm-management
(Zeuzera pyrina L.). surveys are available. On the basis of
Pheromones (sexual attractants) are surveys of farm-management records carried
used to confuse pests and are increasingly out by the FiBL and based on experience it is
used to control codling moth, tortrix moth possible to give the following estimates.
and the moth Laspeyresia lobarzewski. (Even
though these are synthetic products, they
are mostly permitted under organic stan- 22.19.1 Labour input and direct costs
dards because they are: (i) identical to nat- compared with integrated apple production
ural substances; and (ii) they do not get
into direct contact with either plants or The sum total of direct costs for the various
soil.) A prerequisite for the use of production aids (e.g. herbicides versus
pheromones is that orchards are relatively mechanical hoes, systemic fungicides versus
large and are isolated from untreated trees wetting sulphur) is roughly of the same
and that infestation is not too severe. The order.
border area of the plantation has a higher Training, summer and winter pruning,
risk of infestation from females that are fruit thinning, aisle mulching, etc. have not
attracted into the orchard; it can be pro- been listed in Table 22.9, as there is little
tected with additional granulosis virus difference between the two production sys-
applications. tems in the labour input required for these
Methods involving the release of specially activities.
bred beneficials, such as ladybirds for The main extra burden in organic produc-
aphids, are still at the research stage. tion is blossom thinning in April/May. Since
Table 22.9. Labour input: comparison of integrated production (IP) and organic production (from Schmid
and Mouron, 1997).
IP holdings Organic holdings

Activity (man-hours ha1) (man-hours ha1)
Blossom thinning in May 70 70 (annual bearers)

150 (biennial bearers)
Plant protection incl. monitoring 30 (12 applications) 20 (5–9 applications for robust cultivars)
50 (15–20 applications for susceptible cultivars)
In-row management 8 (with herbicide) 25 (mulching, permanent ground cover + area
around tree trunk hand-hoed)
45 (6 times for hoeing, once for cleaning
around trunk)
Control of voles + mice 5 (poisoned bait) 10–20 (traps, CO2 fumigation)
Fertilizing 2 (once per year) 5 (once per year + composted farmyard
manure every 5 years)
Harvest 300 270 (c. 15% lower yield but smaller fruit reduce
harvesting efficiency)
Grading/sizing 90 90 (lower yield and more tolerant grading rules,
but more time-consuming because of higher
number of marred fruit and limited mechanical
sorting equipment)
Additional labour in organic production:
minimum 0 man-hours, maximum 125 man-hours
Apples - Ch.22 21/3/03 3:03 pm Page 581
during this period the plant-protection mea- mechanical blossom-thinning tools are as
sures required are also more labour-intensive yet not very widespread);
(and crucial for success), this is a time of ● smaller proportion of high-yielding
above-normal peak labour demand, particu- cultivars, such as ‘Golden Delicious’ and
larly on holdings also with livestock enter- ‘Jonagold’;
prises and forage cropping. ● more plant-protection problems;
● fewer options for short-term corrections
of nutrient deficiencies;
22.19.2 Yields ● ground-cover vegetation has a greater
impact on the trees’ performance (compe-
Depending on the individual situation tition for nutrients and water), control is
(location, cultivars, planting system) and the labour- and cost-intensive;
expertise of the farm manager, apple yields ● generally fewer trees per hectare (particu-
vary substantially. Surveys conducted dur- larly in older orchards).
ing the past few years have indicated
The more tolerant grading rules for
10–30% lower total yields than conventional
organic production generally allow for a
or integrated production orchards at the
higher proportion of dessert-quality fruit. In
same level of production intensity.
combination with higher prices, strong
The reasons for the lower total yields
demand from distribution channels and, in
were:
certain cases, public-support payments (e.g.
● larger problems with biennial bearing conversion subsidies or agro-environmental
(manual methods are much more payments), the lower yield levels and the
labour-intensive than chemical thinning; higher labour costs can thus be compensated.
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Pflanzgut im ökologischen Obstbau. Bioland Bundesverband, Mainz, 4 pp.
Bertschinger, L. and Stadler, W. (1998) Mechanische Ausdünnung – von Apfelanlagen mit dem Fadengerät.
Leaflet 401, Swiss Federal Research Station for Fruit Wine and Vegetable Growing, Wädenswil, 4 pp.
Bertschinger, L., Stadler, W., Weibel, F.P. and Schumacher, R. (1998) New methods for an environmentally
safe regulation of flower and fruit set and of alternate bearing of the apple crop. In: Proceedings
Second Workshop on Pome Fruit Quality. Bonn-Röttingen, Germany, pp. 64–70.
BIO-SUISSE (1999) Richtlinien für die Erzeugung, Verarbeitung und den Handel von Produkten aus biologis-
chem (ökologischem) Anbau. BIO-SUISSE, Missionsstr. 60, CH-4055 Basel. (Fassung 1. Jan. 1999), 102
pp. (Available in an English translation: Standards for the Production, Processing and Marketing of
Produce from Organic (Biological/Ecological) Farming. Downloadable PDF-files of the German
and English versions at http://www.bio-suisse.ch
Brunner, H., Häseli, A. and Weibel, F. (2000) Biologischer Obstbau auf Hochstämmen, 1st edn. Research
Institute of Organic Farming (FiBL), Frick, Switzerland, 24 pp.
Dolega, E., Bertschinger, L., Fankhauser, F. and Stadler, W. (1997) Zur Untersuchung von Obstknospen:
Aussagekraft und Anwendungsmöglichkeiten. Schweizerische Zeitschrift für Obst- und Weinbau 25,
633–636.
EEC (2000) Council Regulation (EEC) No 2092/91 of 24 June 1991 on organic production of agricultural
products and indications referring thereto on agricultural products and foodstuffs. Official Journal L
198, 22/07/1991, 1–15; http://www.prolink.de/~hps/
Faby, R. and Naumann, W. (1986) Einfluss der N-Düngung von Apfelkulturen auf Nitratgehalte im
Boden, stickstoffhaltige Reservestoffe im Baum und Ertrag. Zeitschrift für Pflanzenernährung und
Bodenkunde 149, 639–657.
Faby, R. and Naumann, W.D. (1987) Die Bedeutung der Einlagerung von Reservestoffen im Herbst bei
Apfelbäumen, dargestellt an Entblätterungsversuchen. Erwerbstobstbau 29, 51–56.
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Marketing of Organically Produced Foods, Vol. 1. CAC/GL 032-1999; http://www.codexalimentar-
ius.net/standard/volume1a/vol1a_e.htm
Häseli, A. (1996) Richtlinienvergleich zum biologischen Obstbau in einigen europäischen Staaten.
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Organic Farming (FiBL), Frick, Switzerland, 20 pp.
Häseli, A. and Bosshard, E. (1993) Förderung von Methoden des biologischen Apfelanbaues 1985–1993.
Internal report, Research Institute of Organic Farming (FiBL), Frick, Switzerland, 167 pp.
Häseli, A., Weibel, F. and Brunner, H. (2000) Sortenempfehlung für den biologischen Erwerbsobstbau auf
Hochstämmen. Research Institute of Organic Farming (FiBL), Frick, Switzerland, 24 pp.
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General Assembly, November 1998, Mar del Plata, Argentinia. 67 pp. http://www.ifoam.org/
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sie, was bringt sie? Obstbau–weinbau 35, 219–222.
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Apples - Ch.23 11/4/03 11:02 am Page 585
23 Principles and Practices of

Postharvest Handling and Stress
Christopher B. Watkins
Department of Horticulture, Cornell University, Ithaca, New York, USA

23.2 Fruit Maturity and Ripening 586
23.2.1 Harvest timing and fruit quality 587
23.2.2 Harvest indices 589
23.2.3 Maturity programmes 592
23.3 Harvesting and Handling 593
23.3.1 Harvest management 593
23.3.2 Postharvest treatments 593
23.4 Storage 595
23.4.1 Temperature 595
23.4.2 Relative humidity 600
23.4.3 Controlled-atmosphere storage 600
23.4.4 Stress treatments 604
23.5 Physiological and Pathological Disorders 604
23.5.1 Physiological disorders 604
23.5.2 Pathological disorders 607
23.1 Introduction changes that contribute to fruit acceptability

occur after harvest, e.g. conversion of starch
The term ‘postharvest handling’ encapsulates to sugars. However, postharvest handling is
the many management decisions and largely concerned with maintaining, rather
processes that are involved in harvesting, than improving, quality. Consequently, the
handling, storage, packaging and transport of management tools used by apple growers,
apple fruit necessary to provide the consumer storage operators, packers and shippers are
with an acceptable product. Most characteris- focused on the following:
tics that influence acceptance of fruit in the
market-place are present at the time of har- 1. Reducing metabolic rates of processes that
vest, and include size and shape, colour and result in undesirable changes in colour, com-
freedom from blemishes (Plate 23.1). Internal position, texture, flavour and nutritional status.
characteristics at harvest include the presence 2. Reducing water loss that can result in loss
of or the potential to develop acceptable of marketable weight, shrivelling, softening
varietal flavour and texture. Some important and loss of crispness.
Apples - Ch.23 11/4/03 11:02 am Page 586
586 C.B. Watkins
3. Minimizing bruising, friction damage and agement through effects on ripening rates
other mechanical injuries. and physiology. Early-maturing apple culti-
4. Preventing the development of physio- vars usually produce much higher levels of
logical and pathological disorders. ethylene than later-maturing ones and have
the shortest storage life. Cultivars with lower
These objectives are met by harvesting the
ethylene-production rates generally have
fruit at the maturity stage that will meet
longer storage potential (Watkins et al.,
the quality standards required by the market
1989b; Gussman et al., 1993; Sunako et al.,
either at harvest time or after storage, by
1999). Responses of fruit to storage environ-
handling fruit carefully and rapidly to avoid
ments are influenced by physiological fea-
mechanical injury and to minimize deterio-
tures, such as diffusion characteristics of the
ration and by applying protective chemical
skin. ‘Golden Delicious’, for example, tends
preservatives (antioxidants, fungicides).
to shrivel faster than other cultivars because
Fruit should be cooled quickly to remove
of breaks in the cuticle, and the Marshall
field heat, and proper refrigeration should
be applied during storage. High relative ‘McIntosh’ strain is less tolerant to low oxy-
humidity should be maintained and the gen in controlled-atmosphere (CA) storage
storage atmosphere should be controlled. than other ‘McIntosh’ strains because of
Protective containers and packaging should higher resistance of the skin to gas exchange
be used during transport and marketing, (Park et al., 1993). Orchard management
along with refrigeration. Good sanitation practices and climatic conditions have an
practices must also be maintained through- impact not only on fruit quality at harvest
out these processes, as well as during har- but also on tolerances of fruit to postharvest
vesting, as outbreaks of food-borne illness storage conditions. Examples of the impacts
can have a devastating effect in the market. of preharvest factors are provided in this
The harvesting of any horticultural prod- chapter, but the reader is referred to Sharples
uct involves the imposition of stress – that is, (1973), Bramlage (1993) and Ferguson and
an interruption, restriction or acceleration of Boyd (2002) for greater detail.
normal metabolic processes. However, many
recommended postharvest conditions also
cause stress, making the term in a posthar- 23.2 Fruit Maturity and Ripening
vest context somewhat nebulous (Kays,
1997). During postharvest storage and han- The apple is classified as a climacteric fruit
dling, stress is an external factor that will (Kidd and West, 1924). The term ‘respiratory
result in undesirable changes in quality if the climacteric’ describes the rise in respiration
fruit is exposed to it for a sufficient duration rate that accompanies the maturation and
or at a sufficient intensity. However, while ripening phase in apple, and this rise is
recommended storage conditions for apple associated with increases in internal con-
represent stress to the fruit, to the posthar- centrations of carbon dioxide and ethylene,
vest physiologist they represent the opti- respiration and autocatalytic ethylene produc-
mum conditions for maintenance of product tion (Fig. 23.1). Climacteric fruit can be sepa-
quality (Kays, 1997). rated from non-climacteric fruit by responses
The objective of this chapter is to outline of respiration and/or ethylene production to
the principles and practices involved in exogenous ethylene or its analogues, such as
postharvest handling and stress. For general propylene (Wills et al., 1998). In a climacteric
postharvest principles, the reader is referred fruit such as the apple, ethylene advances the
to Kader (1992), Kays (1997) and Wills et al. timing of the climacteric, autocatalytic produc-
(1998). The effects of cultivar and preharvest tion continues after removal of ethylene and,
management will not be addressed in detail in contrast to a non-climacteric fruit, the mag-
here, but it should be recognized that the nitude of the respiratory rise is independent of
responses of fruit to postharvest treatments the concentration of applied ethylene. Thus
are greatly affected by these factors. Cultivar, timing of the climacteric and ripening of apple
for instance, can influence postharvest man- fruit is advanced by exposure to ethylene.
Apples - Ch.23 11/4/03 11:02 am Page 587
Principles and Practices of Postharvest Handling 587
Log ethylene production

2
3
Log internal ethylene

concentration (µl l–1)
(µl 10 kg–1 h–1)
1
2
0
1
–1
0
–2
–1
120
4
Respiration (ml 10 kg–1 h–1)

Internal carbon dioxide
10
concentration (%)
3
100
2 90
80
70
60
0 –5 0 5 10
Time from reference time (days)
Fig. 23.1. Patterns of ethylene and carbon dioxide concentrations and production in ripening apple fruit
(from Reid et al., 1973). Respiration (); internal carbon dioxide concentration (); ethylene production ();
internal ethylene concentration ().
Ripening of apple fruit involves many ripening period has a profound effect on the
physiological and biochemical changes. storage quality of fruit. As quality factors
From an applied perspective, the most such as flavour and aroma of the fruit
important of these are softening, the change increase, the storage potential of the fruit
of background or ground colour from green decreases (Fig. 23.2) and therefore harvest
to yellow, loss of acidity, conversion of starch decisions are a compromise between the
to sugars, formation of cuticular waxes and quality and storability of fruit. The increases
synthesis of aromatic compounds. The in fruit quality are typically associated with
metabolism involved in these changes has the climacteric. The length of storage of
been described by Knee (1993). In the natural apples can usually be increased by harvesting
world, these changes result in a desirable fruit before they are fully mature, but quality
product for seed dispersal by the activity of characteristics such as colour and varietal
animals and/or breakdown and decay of the flavour develop less in these fruit, and they
fruit. Many of these changes are at least can be more susceptible to physiological dis-
partly desirable for human consumption, orders, such as bitter pit and superficial
and the objective of apple industries is to scald. Fruit harvested over-mature tend to be
harvest fruit at the appropriate maturity and softer and more easily damaged and may
apply postharvest technologies to control the have water-core and be more susceptible to
rates of these changes in order to provide the diseases and physiological disorders, such as
consumer with an acceptable product. senescent breakdown.
Superimposed on the relationship
between quality and storability during
23.2.1 Harvest timing and fruit quality ripening is that between the harvest period
and acceptable storage length. The harvest
The harvest date within the maturation and period over which market quality of the fruit
Apples - Ch.23 11/4/03 11:02 am Page 588
588 C.B. Watkins
100 harvest periods involved in Fig. 23.3 may be

affected by cultivar. For example, the opti-
mum harvest period for an early-season apple,
Storability Quality
75 (firmness, acidity, (colour, flavour,
such as ‘McIntosh’, is much shorter than that
Relative change
starch, background sugar/acid ratio, for a late-harvested apple, such as ‘Rome

colour)
starch index) Beauty’. Therefore, the x axis in Fig. 23.3 might
50 be 1–2 weeks for the former cultivar and 3–4
weeks for the latter one. Regional differences
can also be important. For example, the ‘Cox’s
25 Orange Pippin’ apple is a short-term storage
Ethylene apple when grown in New Zealand, but a
production long-term storage apple in England.
0 The importance of determining the opti-
0 100
Maturation and ripening period mum harvest date for storability will be
related to how fruit are utilized and there-
Fig. 23.2. Schematic illustration of the increase in fore the expectations of the consumer for the
apple-fruit quality during maturation and ripening product. These expectations vary greatly,
and concomitant loss of storage potential. An depending on the market. Continental
increase in ethylene concentration in the fruit is
Europe, for example, prefers apples at a
generally associated with these changes. (Modified
from Lau, 1985.)
more advanced stage of ripeness than does
the UK (Fidler, 1973). Greater flavour, associ-
ated with tree-ripened fruit, is likely to be a
will be acceptable is relatively wide for fruit premium factor in ‘pick-your-own’ or gate
stored for short periods (1–2 months), but sales during autumn. In contrast, for long-
decreases as storage length increases (Fig. term-stored fruit, texture is more likely to be
23.3). In mature fruit stored for only a month a critical acceptance factor, although accept-
or two, acceptable flavour can develop in able flavour must be present. The require-
early-harvested fruit, while, in those fruit har- ment for a maturity programme generally
vested later, flavour is good and fruit are mar- increases as an industry becomes larger and
ketable if texture is maintained. In contrast, more complex, especially if it is reliant on
the harvest window for a fruit that is mar- long-term storage technology.
keted after long-term CA storage for 9–12 Ethylene production of fruit can be man-
months is much narrower, being limited by ipulated before harvest, either by use of
failure to develop flavour, loss of texture and ethephon application to advance ripening
development of storage disorders. The actual (Larrigaudiere et al., 1996; Stover et al., 2003),
100
Relative fruit quality after storage
Short-term
storage
75
50
Medium-term
storage
25
Long-term
storage
0
0 1 2 3 4 5 6 7 8 9 10
Relative harvest period
Fig. 23.3. Schematic illustration of the effect of harvest period on storability of apples.
Apples - Ch.23 11/4/03 11:02 am Page 589
or by use of aminoethoxyvinylglycine (AVG), of these are indicators of quality, rather than

available commercially as ReTain®, to reduce maturity per se. Moreover, harvest decisions
ethylene production and delay ripening have to be based not only on physiological
(Autio and Bramlage, 1982; Stover et al., 2003). maturity, but also on market requirements,
Harvest dates must be adjusted accordingly. which include factors such as blush, back-
ground colour and usually the absence of
water-core. Therefore, the term ‘harvest
23.2.2 Harvest indices indices’ is a more accurate terminology for
the factors used in making harvest decisions.
Many methods have been proposed as in- The most important harvest indices are as
dices of apple maturity (Table 23.1). Several follows.
Table 23.1. Examples of methods evaluated as indices for determining harvest dates of apple fruit.
Harvest index Referencea
Ethylene production or internal ethylene Dilley, 1980; Lau, 1985; Blanpied, 1986; Chu, 1988;
concentration Knee et al., 1989
Respiration Wilkinson and Sharples, 1967; Blanpied, 1969;
Knee et al., 1989
Starch pattern index Reid et al., 1982; Knee et al., 1989; Blanpied and Silsby,
1992
Starch content Knee et al., 1989
Firmness Lau, 1985; Knee et al., 1989
Soluble-solids concentration Lau, 1985
Firmness and soluble-solids concentration Blanpied, 1974
(sliding scale)
Firmness/soluble-solids concentration Streif, 1983; DeLong et al., 1999
starch index
Background colour Lau, 1985; Watkins et al., 1993; Plotto et al., 1995
Calendar date Blanpied, 1964; Fidler, 1973
Days from full bloom Haller, 1942; Truter and Hurndall, 1988
Heat-unit accumulation Eggert, 1960
Days from full bloom and temperature records Luton and Hamer, 1983; Blanpied and Silsby, 1992;
Beaudry et al., 1993
Water-core incidence Lau, 1985; Blanpied and Silsby, 1992
‘T stage’ Stoll, 1968; Faragher et al., 1984
Flesh colour Lau, 1985
Seed colour Lau, 1985
Free sugars and phosphorylated intermediates Knee et al., 1989
of carbohydrate metabolism
Enzymes (carboxylic ester hydrolase, Knee et al., 1989
NADP malic enzyme, ß-galactosidase)
Peel pigments (chlorophyll, xanthophyll Knee et al., 1989
esters, carotenoids)
NADPH fluorescence Cavalieri et al., 1988
Fruit size Lau, 1985
Loss of bitter flavour Blanpied and Silsby, 1992
Separation force Smock, 1948; Lau, 1985
Titratable acidity Lau, 1985
Visible/near-infrared spectroscopy Peirs et al., 2000
a Referencesare provided as examples only and are not a comprehensive list.
NADP(H), nicotinamide adenine dinucleotide phosphate (hydrogen).
Apples - Ch.23 11/4/03 11:02 am Page 590
590 C.B. Watkins
23.2.2.1 Ethylene production or internal terns, as complications can occur under cer-
ethylene concentration (IEC) tain growing conditions. For example, starch
indices may appear more advanced than
Ethylene, which is an important hormone
other harvest indices because initial starch
that affects fruit ripening, can be measured
accumulation in the fruit was not complete.
relatively easily by gas chromatography.
Therefore, absence of 100% starch staining
Since a rise in ethylene production is associ-
may suggest incorrectly that starch hydro-
ated with initiation of ripening, it has been
lysis has already occurred.
suggested that ethylene production or IEC
should be a major determinant of harvest
decisions (Dilley, 1980; Lau, 1985). However, 23.2.2.3 Flesh firmness
the importance of ethylene in making Apples soften after reaching full size and as
harvest decisions is not straightforward. they mature and ripen (Harker et al., 1997).
Relationships between ethylene production Flesh firmness has been used as a maturity
and optimum harvest dates can be poor index, but it is affected by many preharvest
(Blanpied, 1986; Blankenship and Unrath, factors, including season, orchard location,
1987; Blanpied and Silsby, 1992). Also, the nutrition and exposure to sunlight, which
timing, or presence, of increased ethylene are independent of fruit maturity (Blanpied
production is a function of cultivar and, et al., 1978; Reid et al., 1982). However, as an
within a cultivar, is greatly affected by fac- indicator of internal quality, it can provide
tors such as growing region, orchard within information that is important to fruit perfor-
a region, cultivar strain, growing-season con- mance in storage (Lau, 1985). It can directly
ditions and nutrition (Blanpied and Silsby, affect consumer satisfaction (Liu and King,
1992). Ethylene production may not be 1978), and firmness is being used as a quality
relevant for determining the harvest of criterion by wholesalers, especially in
some cultivars, such as ‘Golden Delicious’, Europe, rather than as a maturity index.
because it does not increase during the
harvest period (Lau, 1985; Chu, 1988; Watkins
et al., 1989b). Ethylene production may be a 23.2.2.4 Soluble-solids concentration
better indicator of when to complete the The soluble-solids concentration of apples
harvest, especially in cultivars such as generally increases as fruit mature and ripen,
‘McIntosh’, where autocatalytic ethylene principally by the conversion of starch to
production precedes preharvest drop sugars (Brookfield et al., 1997). It is also a
(Blanpied and Silsby, 1992). quality index, rather than a maturity index,
being affected by many preharvest factors,
23.2.2.2 Starch test and levels do not necessarily reflect fruit
maturity (Reid et al., 1982; Lau, 1985). As
The hydrolysis of starch to sugars as fruit with firmness, soluble-solids concentrations
ripen can be estimated by staining starch are increasingly being used as a quality crite-
with iodine solution. The resulting patterns, rion by wholesalers. Acceptability of apples
which reflect the extent of starch hydrolysis, may be affected by an interaction of firmness
are rated numerically, using starch charts. and soluble-solids concentration, e.g. a softer
Many starch charts are available, either fruit with high soluble-solids concentration
specific to cultivar (Reid et al., 1982; Lau, may be acceptable, whereas the same fruit
1985), or generic (Fig. 23.4). The starch test with low soluble solids is not. Sliding scales
has become popular because of its ease of incorporating both factors have been devel-
use. Optimum starch indices are available oped (Blanpied, 1974).
for many cultivars and, when the change of
indices is linear, the test can be used to pre-
23.2.2.5 Titratable acidity
dict optimum harvest dates. Indices devel-
oped for a cultivar in one region cannot Malic acid is the predominant acid contribut-
necessarily be applied to other regions. Also, ing to the titratable acidity (TA) of apple
caution is required in interpreting starch pat- fruit. TA decreases during maturation and
Apples - Ch.23
11/4/03
11:02 am
Principles and Practices of Postharvest Handling
Page 591
1 2 3 4 5 6 7 8
100% 50% 0%
Starch–iodine index
Core stain
100% 80% 60% 40% 20% 0%
Flesh stain
Fig. 23.4. Generic starch–iodine index chart (from Blanpied and Silsby, 1992).
591
Apples - Ch.23 11/4/03 11:02 am Page 592
592 C.B. Watkins
ripening, but optimum values vary by culti- flesh-colour changes are used to determine
var and season (Lau, 1985). harvest dates for ‘Tydeman’s Red’ and
‘Spartan’ (Lau, 1985).
23.2.2.6 Background or ground colour
As fruit mature and ripen, the background 23.2.3 Maturity programmes
colour changes from green to yellow, reflect-
ing the loss of chlorophyll. Ground colour Traditionally, maturity programmes have
has been used as a maturity indicator focused on minimizing wastage resulting
(Smock, 1948; Lau, 1985), but early harvest of from physiological disorders and rotting.
fruit for long-term storage may pre-empt the However, better attention to nutrition and
colour changes. Preharvest factors, especially the use of fungicides has reduced the impact
those that affect nitrogen content, can of these factors, and eating quality should
markedly influence chlorophyll concentra- now be a major factor in harvest decisions.
tions, independent of maturity changes (Lau, Apple-growing regions vary in the type
1985). Also, in apples it is sometimes difficult and extent of maturity programmes. IEC
to separate the initial decline of chlorophyll measurements and the starch index have
on a surface area resulting from fruit expan- become the most widely used maturity
sion from the subsequent loss resulting from indices. Background colour is considered an
chlorophyll breakdown (Knee et al., 1989). important harvest index for some bicoloured
Nevertheless, assessment of background apples. In some cases, state regulations have
colour is used commonly for bicoloured been established to set minimum harvest
cultivars, such as ‘Gala’, ‘Braeburn’ and maturities, e.g. the starch index for ‘Granny
‘Fuji’ (Watkins et al., 1993; Plotto et al., 1995). Smith’ in California. Firmness and soluble-
Commercially, colour is usually assessed with solids contents, while not maturity indices
colour charts, but it is commonly measured per se, are often measured at harvest, as they
experimentally using chromameters. can indicate whether fruit will have the mar-
ket quality required by importing countries
such as the UK.
23.2.2.7 Calendar date, days from full bloom,
In the USA, New York and Michigan
and temperature records
operate apple-maturity programmes in con-
The usefulness of full-bloom dates and days junction with their Land Grant universities.
after full bloom, with and without tempera- Currently in Washington State, individual
ture records, varies greatly by cultivar and packing-houses conduct their own maturity
growing region (Luton and Hamer, 1983). programmes in line with their marketing
Calendar dates alone have limited value in strategies. In each region, a wide range of
regions where temperature variations result maturity and quality indices are collected
in wide differences in bloom dates, but, in and the optimum harvest period, sometimes
more consistent growing regions, days from called the harvest window, is established for
full bloom can be the most reliable harvest each cultivar (Anon., 1986; Blanpied and
index for several cultivars (Blanpied, 1964; Silsby, 1992; Beaudry et al., 1993). The Streif
Fidler, 1973). The temperatures during the index (firmness/soluble-solids concentration
first 4–6 weeks after full bloom have been starch index), developed by Streif (1983),
useful in predicting a harvest window for has shown potential as an indicator for
‘McIntosh’ (Blanpied and Silsby, 1992; concluding the harvest of several cultivars
Beaudry et al., 1993). for CA storage (DeLong et al., 1999).
Of the many other indices that have been Some programmes are based on the use of
tested (Table 23.1), their usefulness as har- absolute maturity indices, but factors associ-
vest indices has been variable, depending on ated with marketing cannot be ignored. It
the cultivars and growing regions in which can be argued that the strength of a maturity
they have been utilized. Seed-colour change, programme lies not in reliance on absolute
for example, is specific to ‘McIntosh’, while maturity indices, but on discussion with
Apples - Ch.23 11/4/03 11:02 am Page 593
industry personnel on changes in maturity Samim and Banks, 1993). In general, bruise
and quality that are occurring over the har- susceptibility of apples increases with later
vest period. These will include commercial compared with earlier harvest time and
factors, such as colour and susceptibility to decreases during storage. However, the liter-
bruising. In this way, full participation of ature indicates that the effect of each of these
extension personnel, growers and storage factors is inconsistent, suggesting that the
operators can ensure that fruit of appropriate importance of each factor may vary accord-
storage potential are directed towards short-, ing to the particular handling operation.
medium- or long-term storage. For example,
in years where fruit maturity is more
advanced than usual, but acceptable colour
23.3.2 Postharvest treatments
is not present, the rates of harvesting can be
increased in recognition of a smaller harvest
23.3.2.1 Superficial scald inhibitors and
window for long-term storage.
fungicides
Superficial scald is a physiological disorder
23.3. Harvesting and Handling of apples that develops during cold storage
(Section 23.5.1). It was the major cause of
23.3.1 Harvest management fruit loss during storage until the commer-
cial development of diphenylamine (DPA)
Bruising is a major factor in the downgrading as a scald inhibitor. Susceptibility to the
of fruit in the market-place. Most bruising disorder can be affected by cultivar and can
results from impact damage, although com- decline with later harvest date. Several major
pression bruising, which occurs when exces- cultivars, such as ‘Delicious’ and ‘Granny
sive weight is placed on fruit in containers or Smith’, are highly susceptible during normal
when cartons are damaged, can cause fruit commercial harvest periods.
loss (Plate 23.2). Impact injury can occur DPA is typically applied as a drench, often
during all facets of fruit handling from at the storage receiving point while bins of
harvest to the consumer. These include the fruit are still on trucks, or by bin dipping.
following: The concentrations necessary for scald con-
trol vary by cultivar, and label rates on
1. The harvest operation, where bruising can
proprietary DPA products from commercial
be reduced by training of pickers, using
suppliers should be followed. Cultivars also
appropriate picking bags and bins and care-
vary in sensitivity to DPA-induced skin
ful transport of bins from the field to the
injury. Several factors that affect DPA efficacy
storage or packing-house.
have been summarized by Little and Holmes
2. During the packing operation, impacts
(2000). Lower DPA application rates may be
can occur by fruit-to-fruit contact or by
used if scald risk is lower, for example,
impacts with the grading equipment.
because of climate. DPA use can be avoided
3. Transport of packed cartons of fruit to the
where cultivars, such as ‘Empire’ and ‘Gala’,
market-place and handling of fruit at the
have a low scald risk.
retail level (Plates 23.2 and 23.3).
The use of DPA is widespread, although it
It has been estimated that 40% of bruising is not registered for use in some countries.
occurs in the field, 40% during grading and Adequate control of scald may be obtained
20% during transport (Funt et al., 1999). by rapid cooling and rapid CA storage, espe-
Many factors that affect apple-fruit suscepti- cially low oxygen concentrations, but effi-
bility to bruising have been investigated. cacy can vary according to cultivar and
These have included the effects of orchard- growing region. Alternative means of con-
management systems (Baugher et al., 1996), trolling scald development, for example,
but most focus has been on cultivar, fruit stress treatments (Section 23.4.4), have also
size, harvest date, storage periods, tempera- been investigated because of concern that
ture and turgor of the fruit (Klein, 1987; DPA may become unavailable for this pur-
Apples - Ch.23 11/4/03 11:02 am Page 594
594 C.B. Watkins
pose due to consumer concerns about food et al., 1998). The detergent wash before wax
safety relating to the use of such chemicals. application can also remove dust, dirt and
An additional issue with the use of DPA contamination arising from washing in dirty
treatment is that fungicides are also applied water, which can occur in non-chlorinated
in the solution to reduce fungal decay. hydrohandlers (Little and Holmes, 2000).
Fungicides may also be subject to consumer
concerns about food safety. Development of
23.3.2.4 1-Methylcyclopropene (1-MCP)
pathogen resistance to benzimidazole fungi-
cides is discussed in Section 23.5.2. Controlling ethylene production and action
is a primary goal in the postharvest manage-
ment of apples, and currently available
23.3.2.2 Calcium
methods of achieving this are discussed in
Treatment of fruit with calcium can reduce Section 23.4. However, a new compound,
development of bitter pit, senescent break- 1-MCP, is now available for many apple
down and rots (Wilkinson and Fidler, 1973; industries, under the commercial name
Sharples and Johnson, 1976), and may slow SmartFreshTM, Registration for 1-MCP was
softening (Poovaiah et al., 1988; Saftner et al., obtained in the USA and other countries in
1998). Depending on growing region and 2002 and European registration is pending.
cultivar, storage operators will therefore 1-MCP is structurally related to ethylene, has
apply calcium, usually as food-grade flake a non-toxic mode of action and is applied at
calcium chloride or proprietary formulations. very low dose levels, with low measurable
Cultivars vary in susceptibility to calcium- residues in food commodities. The US
induced skin burn, and product-label appli- Environmental Protection Agency has classi-
cation rates should be followed to minimize fied 1-MCP as a plant growth regulator.
risk. 1-MCP is thought to bind to the ethylene
Fruit should be treated with calcium receptor, competing with ethylene for the
immediately after harvest, and it is applied available binding sites and thereby prevent-
either alone or with DPA (and sometimes ing ethylene action (Sisler and Serek, 1997).
with a fungicide). Application is usually by Whereas ethylene diffuses rapidly from the
drenching or dipping. Calcium treatment is binding site after ethylene treatment, this
only commercially effective where disorder compound remains bound for long periods
incidence is low to moderate, but effective- and formation of an active complex is
ness may be improved by CA storage prevented. Inhibition of ethylene action may
(Ferguson and Watkins, 1989). Factors affect- eventually be overcome by the production of
ing use of postharvest calcium treatments are new receptors (Sisler et al., 1996).
discussed by Little and Holmes (2000). Softening, yellowing, respiration, loss of
TA and sometimes a reduction in soluble-
solid concentration, as well as the develop-
23.3.2.3 Waxes
ment of several physiological disorders, are
Apples are commonly waxed with shellac- delayed or inhibited by 1-MCP application
or carnauba-shellac-based coatings during (Fan et al., 1999a,b; Rupasinghe et al., 2000;
the packing-line operation, although some Watkins et al., 2000). Responses of fruit to 1-
markets prefer unwaxed fruit. Also, because MCP may be affected by cultivar and fruit
shellac and carnauba-shellac are associated maturity. Volatile production by apples is
with non-food uses, such as floor and car also inhibited by 1-MCP (Fan and Mattheis,
waxes, alternative coating materials are 1999; Rupasinghe et al., 2000). These results
being tested (Alleyne and Hagenmaier, are consistent with the view that volatile pro-
2000). duction is regulated by ethylene, and con-
The main advantage of waxing is to sumer studies on the acceptability of
improve fruit appearance. Wax coatings can 1-MCP-treated fruit will be required to
extend shelf life of fruit, by reducing water ensure that flavour is not unacceptably
loss, respiration rates and ripening (Saftner compromised.
Apples - Ch.23 11/4/03 11:02 am Page 595
23.4 Storage transport and retail display. This combina-

tion of events is sometimes described as
Depending on cultivar and growing region, the ‘cold chain’, highlighting the importance
some fruit are stored and marketed for up to of maintaining the links from harvest to
a year, when the following year’s crop consumer.
becomes available. Therefore, unless fruit are
sold for immediate consumption, the major
23.4.1.1 Effects of temperature on metabolism
objective of postharvest management is to
decrease the rates of metabolism and thereby RESPIRATION The respiration rate of fruit is
the rates of deterioration and quality loss. directly affected by temperature (Fig. 23.5)
Commonly, respiratory activity of harvested and the respiratory climacteric is sup-
apples is used as an indicator of metabolic pressed by storage temperatures below
rates. The principal mechanisms available to 10°C (Fidler and North, 1967). However,
slow down metabolism are temperature and respiration of chilling-injury-susceptible
control of storage atmospheres. Control of fruit may be stimulated by colder storage
relative humidity is also an important com- temperatures, e.g. ‘Idared’ fruit stored at
ponent. The interaction of the three factors 0°C initially had lower respiration rates
must be considered in developing a storage than those stored at 2°C and 4°C but the
regime for any cultivar. respiration rate of these fruit increased as
low-temperature breakdown developed
(Johnson and Ertan, 1983). This effect was
23.4.1 Temperature exacerbated by lowered oxygen concentra-
tions in the atmosphere (Table 23.2).
Generally, low respiration rates and longer Increased risk of chilling injuries and of the
storage periods of apples are directly related development of alcoholic off-flavours forms
to lowered storage temperatures. The lowest the basis of recommendations for raising
temperatures for storage must be above storage temperatures when oxygen con-
freezing and those at which chilling injury centrations are lowered (Table 23.3).
will develop. Maximizing quality mainte-
nance of fruit requires attention to tempera- ETHYLENE Low storage temperatures can
ture not only immediately after harvest and slow down the onset of ethylene production
during storage, but also during packing, in apples, but chilling temperatures can also
Respiration rate (mg carbon dioxide kg–1 h–1)
35
30
25
20
15
Summer
10
Autumn
5
0
0 5 10 15 20 25
Temperature (C)
Fig. 23.5. Effect of temperature on respiration rate of summer (early ripening) and autumn (late ripening)
apples (modified from Hardenburg et al., 1986).
Apples - Ch.23 11/4/03 11:02 am Page 596
596 C.B. Watkins
Table 23.2. Respiration rate and incidence of low-temperature breakdown on

‘Idared’ stored at 1, 2 or 21% oxygen at 0, 2 or 4°C (modified from Johnson
and Ertan, 1983).
Oxygen concentration (%)
Storage temperature (°C) 1 2 21
Respiration rate (ml 10 kg1 h1)

0 17 18 36
2 11 10 33
4 9 12 35
SED 1.6
Low-temperature breakdown (%)

0 73 98 78
2 5 0 0
4 0 0 0
SED 5.1
Table 23.3. Ranges of atmospheric and temperature conditions for selected apple cultivars (modified
from Kupferman, 1997; Watkins et al., 2003). References should be consulted for specific atmosphere
and temperature combinations for each growing region.
Oxygen Carbon dioxide Temperature

Cultivar (%) (%) (°C)
‘Braeburn’ 1–3 < 0.5–1.2 0–1.5

‘Bramley’s Seedling’ 1 5 4
‘Cortland’ 2–3 2–3 0
2–3 2–3 for 1 month, 2
then 5
‘Cox’s Orange Pippin’ 1.2–2 0.7–4 3–4
‘Delicious’ 0.7–2.5 0–4.5 0–1.1
‘Elstar’ 1.5–2.5 1–4.5 0–1.5
‘Empire’ 1.5–3 0.5–3a 1–3
‘Fuji’ 0.7–2.5 1–2a 0–1
‘Gala’ 1–3 2–3 0–3
‘Golden Delicious’ 1–3 2–3 0.5–2
‘Granny Smith’ 0.8–2.5 0.5 0.5–2
‘Idared’ 1.3–3 0.5–4 0–4
‘Jonagold’ 1–2.5 2–3 0–3
‘McIntosh’ 3 2–3 for 1 month, 2
then 5
2 2–3 for 1 month, 3
then 5
‘Marshall McIntosh’ 4–4.5 2–3 for 1 month, 2
then 5
‘Mutsu’ 1.5–2.5 1–3 0–1
‘Pink Lady®’ 1.5–2 1 0
a CO
2-sensitive so keep CO2 well below the O2 level. If not treated with DPA, use 1.5–2% CO2 during first
30 days.
Apples - Ch.23 11/4/03 11:02 am Page 597
enhance ethylene production in some culti- with high nitrogen and low calcium contents.
vars (Knee et al., 1983). Accumulation of 1- Rapid cooling appears to be more critical for
aminocyclopropane carboxylic acid (ACC), a fruit that are more likely to ripen quickly.
key intermediate in the ethylene biosynthetic Also, the negative effects of slow cooling on
pathway, and ethylene production of firmness and colour are magnified as storage
‘Granny Smith’ apples at 20°C are markedly length increases. Therefore, inadequate
increased by exposure to 0°C for at least investment of resources at harvest to ensure
8 days, but this does not occur in ‘Royal rapid fruit cooling may not be apparent until
Gala’ or ‘Delicious’ (Jobling et al., 1991; late in the storage period, when fruit may
Larrigaudiere et al., 1997). Cold treatment not meet minimum firmness standards for
appears to reduce resistance to ripening in marketing.
‘Granny Smith’, and the ripening physiology The predominant method of cooling
of this cultivar may be more analogous to apple fruit involves exposing fruit to normal
that of winter pears. air flow in a refrigerated room. However,
this method is slow and inefficient because
CHILLING INJURIES Some fruit types, typically air flows around, rather than through, bins
tropical or subtropical in origin, develop or cartons of fruit. Slow cooling can result in
chilling injury when exposed to storage tem- loss of quality when rooms are filled rapidly
peratures below 10–15°C (Saltveit and and refrigeration capacity cannot cope with
Morris, 1990). Temperate fruit, though resis- large fruit loads. The conversion of many
tant to chilling injury, are not immune to it industries to rapid CA storage (Section
(Bramlage and Meir, 1990). Most apple culti- 23.4.3.3) has also resulted in increased
vars are not sensitive to chilling tempera- pressure to improve fruit cooling rates.
tures and maximum storage life is obtained Faster room cooling of fruit can be
by storing them as close to 0°C as possible. achieved by separating and loading bins or
However, some cultivars are susceptible to cartons into separate rooms for pre-cooling
the development of chilling-related disorders before they are moved into the long-term
below 2–4°C. These disorders specifically storage room (modified room cooling) or by
include low-temperature-related disorders, loading only the quantity that can be
such as low-temperature breakdown and handled by the existing refrigeration system.
soft scald, as well as disorders that are aggra- Modified room cooling involves increased
vated by such temperatures. These disorders movement of fruit and logistical planning,
are discussed in Section 23.5.1. but can be extremely useful early in the har-
vest period when harvest of the faster-respir-
ing early-season apples coincides with the
23.4.1.2 Cooling after harvest
availability of still-empty storage rooms that
Because the respiration rate of apple fruit is will be used later. In such situations, where
affected by temperature (Fig. 23.5), the rate fruit volumes exceed the refrigeration capac-
of fruit cooling after harvest can markedly ity of the cold storage, improved fruit cool-
influence quality maintenance. Fruit should ing can be achieved by placing no more than
be removed from exposure to radiant energy two stacks of bins across the width of each
in the orchard to refrigeration, or at least CA storage room each day (Bartsch and
shade, as quickly as possible after harvest. Blanpied, 1990). In cases where the room is
However, the importance of rapid cooling not down to 0°C by the next morning, the
varies depending on cultivar, harvest number of stacks should be reduced even
maturity, nutritional status of the fruit and further to one-deep. Faster cooling will be
potential storage performance. Early-season obtained if bins are placed in the down-
cultivars tend to soften more rapidly than stream discharge of the evaporator with pal-
those that mature in the later part of the let runners orientated in the same direction
harvest season. Within a cultivar, apples tend as the air flow. Additional bins of fruit
to soften more rapidly at later stages of should be stacked, no more than two-high,
maturity than at earlier stages and in fruit in unfilled refrigeration rooms to cool
Apples - Ch.23 11/4/03 11:02 am Page 598
598 C.B. Watkins
overnight before loading into the CA room Additional factors in selecting storage tem-
the next morning. These stacks should be peratures include the interaction between
placed randomly throughout the unfilled temperature and low oxygen and/or high
room to maximize air exchange with the carbon dioxide concentration (see Section
fruit. The capacity to cool fruit is dependent 23.4.3.3). Also, temperature affects humidity
upon the refrigeration capacity and room requirements – it is easier to maintain rela-
design. tive humidity above 90% at 1°C than at 0°C.
Forced-air cooling and hydrocooling are Temperature should be monitored through-
less commonly used than room cooling for out the storage period by measurements at
apples, although forced-air cooling is being different points throughout the storage room
used increasingly for cooling of fruit in with thermocouples (Bartsch and Blanpied,
cartons. In forced-air cooling, bins or cartons 1990). It is dangerous to rely on a single ther-
are stacked in patterns so that cooling air is mometer at the door, as temperatures within
forced through, rather than around, the indi- stacks and throughout the room may be
vidual container. A slight pressure gradient, lower or higher than indicated by such
usually developed with a fan, forces cold air readings.
through the containers, which contain vent- Stepwise change of temperature – for
holes placed in the direction that the air will instance, where fruit are kept at 2°C for
move. Also, minimal packing material that 4 weeks, 1°C for the next 4 weeks and finally
will interfere with free movement of air at 0°C, may reduce low-temperature injury
through the container is used. Direct contact and enhance flavour, compared with fruit
of cold air with the product can cool the fruit directly cooled to 0°C. However, storage life
0.1–0.25 times faster than room cooling. The under such regimes may be reduced (Little
cooling rate is controlled by the volume and Holmes, 2000).
of air passing over the product, and the
economics of cooling are affected by the fan
23.4.1.4 Post-storage temperature control
speed and the number of containers being
cooled. Water loss from the product is less While fruit are kept at optimum tempera-
than that found for room cooling. tures during storage, the cold chain is most
In hydrocooling, apple temperatures are frequently broken after storage during pack-
reduced by cold water flowing over the fruit ing, transport and retail display. When stor-
surface, by either flooding, spraying or age rooms are opened for marketing, fruit
immersion of the fruit in chilled water. The can be exposed for extended periods to tem-
rate of internal cooling is related to the size peratures above optimal due to operations
and shape and the thermal conductivity of required for movement of fruit for packag-
the fruit being cooled. The method is simple, ing. Fruit temperatures can increase during
economical and effective and avoids water this time, as well as during packing. After
loss, although it is obviously limited to bins fruit have been packed into plastic bags or
rather than cartons of apples. trays and placed into cartons, it is important
to remove this heat. Passive cooling may
take several days because cardboard cartons
23.4.1.3 Storage temperatures
are good insulators and offer poor air circu-
The recommended conditions for commer- lation (Fig. 23.6). Moreover, fruit respiration
cial storage of apples are 0.5°C to 4°C contributes to temperature increases within
(Table 23.3), the desired temperature being a cartons. Failure to remove heat accumulated
function of sensitivity to low-temperature- during packing processes may be responsible
associated injuries (Section 23.4.1.1). How- for firmness losses during subsequent
ever, these disorders typically develop over storage and transport and have a negative
long-term storage, and sometimes tempera- impact on fruit quality at the consumer level
tures closer to 0°C are used for short-term (Kupferman, 1994; Watkins, 1999). Forced-air
(1–2 months) storage of chilling sensitive cooling of packed fruit, with modification
cultivars to maximize firmness retention. of carton design to improve air flow over
Apples - Ch.23 11/4/03 11:02 am Page 599
fruit, resulting in faster cooling (Fig. 23.6), is container or other transit vehicle. To mini-
becoming more common (see also Section mize warming of the crop and therefore
23.4.1.2). A downside of using vented car- avoid losing the benefits of cooling, it is nec-
tons, however, is that fruit can also warm up essary to ensure fast and efficient loading.
faster if good refrigeration is not maintained. Under ideal conditions the loading dock and
Transport requirements for apples can load assembly areas are refrigerated. Pallets
range from simple, involving short distances containing cartons should be loaded into
to a nearby local market, to sophisticated, transit vehicles in such a way as to allow an
involving long distances internationally by air flow that is adequate for temperature
land and sea. Most transit vehicles have control.
sufficient refrigeration capacity to maintain, Few retail outlets have display refrigera-
rather than cool, fruit. Thus apples should tion. Moreover, typical displays in US super-
be cooled properly before loading into the markets employ stacking of loose fruit, with
(a)
16
14
Passive cooling
12
Core temperature (C)
10
Middle
8
6
Top
4
2 Bottom
0
0 20 40 60 80 100 120
Time (h)
(b)
14
12
Forced-air cooling
Core temperature (C)
10
8
Middle
6
Top
4 Bottom
0
0 20 40 60 80 100 120
Time (h)
Fig. 23.6. Cooling patterns of apple fruit in cartons within a pallet stack that have been allowed to cool
passively (a), compared with forced-air cooling (b). Data are of core temperatures of ‘Empire’ apples kept in
60 cm 40 cm export cartons in the top, middle and bottom of stacked pallets. (From Watkins, 1999.)
Apples - Ch.23 11/4/03 11:02 am Page 600
600 C.B. Watkins
increased risk of bruising. In the UK, cartons storage is outside the scope of this chapter,
such as the 60 cm 40 cm box are used more but readers are referred to Bartsch and
extensively. These cartons have only two Blanpied (1990) and Bishop (1996).
layers of fruit and are placed directly on
display. As well as reducing the need to
23.4.3.1 Effects of CA on metabolism
handle fruit, more rapid turnover of product
minimizes exposure of fruit to elevated RESPIRATION The metabolic rates of apple
temperatures. fruit are reduced by decreasing oxygen
and/or increasing carbon dioxide concentra-
tions in the storage atmosphere. The classic
23.4.2 Relative humidity studies of Fidler and North (1967, 1968)
demonstrated that low oxygen concentra-
Relative humidity is a major factor in pre- tions reduce oxygen uptake and carbon diox-
venting moisture loss of apples and therefore ide production proportionally, and therefore
maintaining their quality. Relative humidity the respiratory quotient remains constant.
of 90–95% is recommended for apples to pre- High carbon dioxide concentrations reduce
vent shrivel. The major causes of dehydra- carbon dioxide production more than oxy-
tion of fruit in cool stores are the small gen uptake, and the respiratory quotient
surface areas of refrigeration coils and/or therefore declines under these conditions.
frequent defrosting. Therefore, CA rooms Decreasing oxygen concentrations delays the
should have the largest coil size feasible for onset of ethylene production in a ventilated
the room. The number of defrost cycles can system in which low ethylene concentrations
also be reduced to optimize relative humid- in the atmosphere are maintained, but not
ity in the room, and some operators reduce in a product-generated atmosphere where
the oxygen concentration to the minimum ethylene can accumulate in the room (Knee,
safe level and then raise the temperature to 1980). Decreasing oxygen concentrations also
1–2°C to minimize the need to defrost. decrease the rates of chlorophyll loss and
Storage rooms can be outfitted with high- softening (Table 23.4) and the associated
pressure water-vapour systems that can release of soluble cell-wall components, such
add moisture to the room and which are as uronides (Knee, 1980).
suited for operation at temperatures around CA may affect respiratory metabolism via
0°C. The air-distribution system should be effects on glycolysis, fermentative metabo-
designed to prevent condensation of water lism, the tricarboxylic acid (TCA) cycle or the
droplets on fruit in order to prevent decay. electron-transport chain, but the extent to
The use of plastic rather than wooden bins or which respiration is affected directly or
of bin-liners inside wooden bins can also whether other cellular processes whose
help minimize shrivel – for example, with energy demand affects respiratory rates are
‘Golden Delicious’. slowed is uncertain. Severe stresses resulting
from injurious levels of either oxygen or car-
bon dioxide induce accumulation of
23.4.3 Controlled-atmosphere storage acetaldehyde, ethanol and ethyl acetate. In
addition, succinate accumulation is a feature
The apple is the predominant horticultural of fruit treated with high carbon dioxide con-
commodity stored under CA conditions. The centrations (Hulme, 1956; Fernández-Trujillo
objective of CA storage is to lower oxygen et al., 2001), probably because of inhibition of
and increase carbon dioxide concentrations succinate dehydrogenase under these condi-
to levels that will maintain fruit quality tions (Shipway and Bramlage, 1973).
by decreasing respiratory metabolism and
reducing ethylene production and action, ETHYLENE CA storage can also affect ethyl-
but not to levels that induce fermentation or ene biosynthesis and action, and therefore the
other damaging events. The technology effects of CA are not solely due to effects of
involved in establishing and maintaining CA these atmospheres on respiration (reviewed
Apples - Ch.23 11/4/03 11:02 am Page 601
Table 23.4. Oxygen effects on ripening of ‘Cox’s Orange Pippin’ apples (modified from Knee, 1980).
Days to lose:
1 g chlorophyll cm2 0.25 g chlorophyll g1

Oxygen (%) in peel in flesh 9.8 N flesh firmness
21 90a 83a 61a

10 125b 131b 74b
5 141c 135b 74b
2 242d 230c 85c
1 –1 216c 191d
1Apparent increase in chlorophyll.
Values with uncommon letters are significantly different at the 5% level.
by Watkins, 2002). Ethylene production is re- of preclimacteric apple fruit in 2% oxygen,

duced by as much as 50% in a 3% oxygen 2% oxygen plus 5% carbon dioxide or air
atmosphere (Burg and Thimann, 1959). Burg plus 5% carbon dioxide, compared with air
and Burg (1967) suggested that low oxygen alone, have been associated with reduced
concentrations impeded ethylene binding, expression of genes encoding ACC synthase
but the primary effects of low oxygen are and ACC oxidase and of ACC-synthase
likely to be directly on ethylene biosynthesis, activity (Gorny and Kader, 1996a). ACC-
since ACC oxidase, a key enzyme in ethylene oxidase activity in air plus 5% carbon di-
biosynthesis, has an absolute oxygen require- oxide, however, was not different from that
ment for activity (Adams and Yang, 1979; in air alone. Primary effects of carbon
John, 1997). dioxide appeared to be on ACC-synthase
The action of carbon dioxide concentra- transcription and on accumulation of active
tions on ethylene biosynthesis is less clear ACC-oxidase protein. Fungistatic levels of
because: (i) it is difficult to separate the direct 20% carbon dioxide or 0.25% oxygen on
effects of elevated carbon dioxide from those preclimacteric and climacteric apples have
of low oxygen concentrations on ethylene also indicated that ACC-synthase transcript
biosynthesis; (ii) solubilization of carbon abundance and enzyme activity may be a key
dioxide produces H+ and HCO 3 , which may step in the inhibition of ethylene production
affect the pH of the cytoplasm and conse- (Gorny and Kader, 1996b, 1997).
quently enzyme activities (Bown, 1985); and
(iii) maximal activity of ACC oxidase in vitro
23.4.3.2 Factors that affect tolerances of
requires carbon dioxide and it is uncertain to
fruit to CA
what extent enzyme activity is regulated
in vivo by the gas. Burg and Burg (1967) Two factors affect the tolerance of apple fruit
proposed that carbon dioxide displaced to low oxygen and elevated carbon dioxide
ethylene from ethylene receptor sites, but concentrations:
carbon dioxide effects are separable from
1. The resistance of the skin to gas exchange
those associated with ethylene perception
between the atmospheres outside and inside
(de Wild et al., 1999).
the fruit.
In apples, low oxygen concentrations
2. Metabolic sensitivity to the gases.
inhibit only the ACC-to-ethylene step, while
high carbon dioxide also inhibits formation The effect of these factors is illustrated by
of ACC (Li et al., 1983). The fruit have lower comparing the effects of oxygen on ethanol
internal ethylene concentrations and ACC concentration (used as an indicator of fer-
accumulation in a 2.5% oxygen atmosphere mentation) in the fruit of ‘Delicious’ and two
than in air (Lau et al., 1984). Patterns of ‘McIntosh’ strains (Fig. 23.7). The ‘Delicious’
delayed and decreased ethylene production apple is very tolerant to low oxygen, where-
Apples - Ch.23 11/4/03 11:02 am Page 602
602 C.B. Watkins
as the ‘Marshall McIntosh’ accumulates pheres, even for the same cultivar (Table
ethanol at higher external oxygen concentra- 23.3), reflects these factors, as well as differ-
tions. ‘Red Max’ is somewhat intermediate ent storage strategies employed in different
between the others. The skin resistance of the growing regions. Thus, a CA recommenda-
‘Marshall’ strain is much greater than that of tion for a cultivar may not apply every-
the ‘Red Max’ strain, and thus the internal where. Regulations on CA storage, covering
oxygen concentrations in the two strains will both the safe operation and the use of the
be similar at external oxygen concentrations legal definition of ‘controlled atmosphere’
of 2% and 0.8%, respectively (Park et al., for stored apples may exist and differ in
1993). However, differences in tissue sensi- different states or countries. These regula-
tivity to low oxygen concentrations between tions include the rate of establishment of CA
strains also exist, as, even at the same conditions, the maximum concentration of
internal oxygen concentrations, ethanol oxygen permitted and the minimum length
accumulations vary. of time the fruit can be in CA.
The effects of oxygen and carbon dioxide CA technology has undergone major
can interact, with fruit sensitivity to low changes in the last 30 years. Old protocols,
oxygen increasing as carbon dioxide concen- where CA rooms were loaded over 8–10 days
trations increase. Sensitivity to both gases is and where fruit respiration then lowered
increased by lower storage temperatures and oxygen to 2.5–3% over an additional 15–20
is dependent on cultivar and, within culti- days, are obsolete. ‘Rapid CA’, in which
vars, on factors such as harvest date (Park the oxygen concentration in CA rooms is
et al., 1993). Injuries associated with oxygen reduced to less than 2.5% by flushing the
and elevated carbon dioxide are discussed in atmosphere with nitrogen within 4–7 days
Section 23.5.1. from the time of harvest to when CA condi-
tions are applied, is now standard practice
23.4.3.3 Recommendations for CA storage for many industries. Some cultivars must be
cooled before rapid CA to avoid injury.
The recommended gas composition and The use of rapid CA is critical for long-
temperature conditions for CA storage are term (> 6 months) storage. Longer periods
specific to cultivar, growing region and between harvest and CA storage can still
sophistication of the equipment available for result in a superior product to that obtained
monitoring and controlling the atmospheres. in air storage, but the benefits of CA storage
The wide range of recommended atmos- for fruit quality may be lost over time. Even
when rooms are filled over extended peri-
ods, oxygen concentrations in the CA rooms
40 are usually lowered by nitrogen flushing. It
is becoming more common to open rooms
Ethanol (mg 100 g–1)
‘Marshall’
30 ‘RedMax’ briefly to remove some of the fruit for
‘Delicious’ marketing, and nitrogen flushing is used to
20 re-establish atmospheres in resealed rooms.
CA storage regimes fall into one of three
10 categories, depending on the sophistication
of equipment and technology involved.
0 1. Standard CA. Standard CA involves
0.0 0.5 1.0 1.5 2.0 2.5 3.0 application of conservative atmosphere con-
Oxygen during 6 days at 2.2C (%) ditions with a minimum risk of gas-related
injuries (Table 23.3). Carbon dioxide concen-
Fig. 23.7. Ethanol concentrations in the flesh of
‘Marshall McIntosh’, ‘Red Max McIntosh’ and trations are typically in the 2–3% range and
‘Delicious’ apples when kept in different oxygen provide additional quality benefits to those
concentrations at 2.2°C (from Blanpied and obtained by using 2–3% oxygen alone.
Jozwiak, 1993). Control of these atmospheres may be manual
Apples - Ch.23 11/4/03 11:02 am Page 603
by daily reading and adjustment or via potassium permanganate, by oxidation or by

computer-controlled equipment. The margin catalytic burners.
of safety is large enough when oxygen Results of this technology for fruit quality
concentrations are around 2% for fluctuations have been variable, as low-ethylene CA stor-
in gas concentrations found in manually age does not maintain quality if concentra-
adjusted storage rooms not to cause fruit tions of ethylene gas within the fruit cannot
injury. be controlled. Because of the resistance of the
2. Low-oxygen CA storage. In low-oxygen skin to diffusion, it is possible to have high
CA storage, fruit are kept at oxygen con- concentrations of ethylene within the fruit,
centrations below 2% but above the concen- with consequent effects on ripening, even
trations at which fermentation will occur when concentrations in the storage atmos-
(Table 23.3). In Europe, the term ultra-low phere are low. Therefore, the technology is
oxygen (ULO) is used to describe these limited to naturally low-ethylene-producing
atmospheres. The safe oxygen concentration cultivars such as ‘Empire’, and/or to early-
varies by cultivar and growing region. harvested fruit to ensure that only preclimac-
‘Delicious’ apples from British Columbia, teric fruit are stored. Loss of firmness, but
Canada, for example, can be stored safely at not colour, may be delayed by low-ethylene
0.7% oxygen concentrations (Lau, 1997), CA storage (Stow et al., 2000). The return on
allowing control of superficial scald without investment in this technology for maintain-
the use of DPA. Fruit of the same cultivar ing fruit quality has not been adequate.
from other growing regions may show injury However, reduced superficial-scald inci-
when stored at these low oxygen levels (Lau dence by lowering ethylene, even when
et al., 1998). Strains within a cultivar can also concentrations are above 1 l l1, in CA
vary in sensitivity (Lau, 1997). In general, it storage (Knee and Hatfield, 1981) could be
is necessary to increase storage temperatures useful if DPA is no longer available to the
when low-oxygen CA storage is used. The apple industry.
carbon dioxide concentrations under low- CA technologies, as well as associated
oxygen storage are usually much lower than marketing strategies, will continue to
under standard CA, because additional develop. For example, decreasing periods of
benefits of carbon dioxide for firmness are acceptability for air-stored fruit, especially of
not observed. softer cultivars, such as ‘Cortland’ and
Protocols have been developed to reduce ‘McIntosh’, are resulting in increasing use of
the risk of low-oxygen injury development short-term CA. This type of storage is occur-
in regions where susceptibility of fruit to ring even though storage time of fruit under
damage appears high. In the north-eastern CA does not meet regulatory requirements
USA, for example, these include factors such for labelling the fruit as being ‘CA-stored’.
as harvesting fruit early in the harvest CA can also be used during shipping, either
period, avoiding fruit with low seed counts, using modules that maintain atmospheres in
rapidly reducing fruit-core temperatures, ships’ holds or by use of CA-fitted containers
raising storage temperatures, avoiding the (Bishop, 1996). Apples are a relatively low-
use of DPA and using automatic gas-analysis value commodity, however, and the utiliza-
and control equipment to eliminate oxygen tion of new and existing technologies is
fluctuations that might lead to low-oxygen influenced greatly by cost-effectiveness. In
injury (Blanpied, 1990). some regions, sophisticated technologies are
3. Low-ethylene CA storage. Low-ethylene not required to meet market demands,
(< 1 l l1) CA storage has been evaluated as whereas, in others, investment in these tech-
a method for reducing superficial scald, as a nologies is the cost of staying in the business.
safe substitute for low-oxygen CA storage Modified-atmosphere (MA) storage, in
and for retarding flesh softening and other which atmospheres around the fruit are
forms of senescence (Knee and Hatfield, modified passively by polymeric-film bags,
1981; Blanpied, 1990; Stow et al., 2000). has been tested for both transport and
Ethylene is removed by absorption on to wholesale purposes (Hewett and Thompson,
Apples - Ch.23 11/4/03 11:02 am Page 604
604 C.B. Watkins
1989; Geeson et al., 1994; Watkins et al., 1977; Lau and Looney, 1978). Also, early
1997a), but have been used on only a limited research showed that treatment of fruit with
scale commercially. The major impediments 30–60% carbon dioxide inhibited superficial
to their use include additional cost of the scald development (Pieniazek et al., 1946),
films, as well as those associated with the but benefits are limited, especially as risk of
logistical requirements of training the packers, carbon dioxide-induced injury may be high
sealing of bags and preventing damage to (Fernández-Trujillo et al., 2001).
bags.
23.4.4.3 Heat treatments
23.4.4 Stress treatments
Heat treatments, applied either as hot air
(e.g. 38°C for 3–4 days) or as short-term hot
There has been increasing interest in short-
water (e.g. 40–50°C for up to an hour),
term stress treatments, such as low oxygen
inhibit softening of apples and delay devel-
and high carbon dioxide concentrations that
opment of superficial scald during storage
are outside the normal range for storage, and
(reviewed by Lurie, 1998). However, heat
heat, for maintenance of quality and disease
treatments can accelerate the rate of degreen-
control and to meet quarantine require-
ing. While hot-water treatments may be suit-
ments. This research has been prompted,
able for quarantine treatments, commercial
for example, by the search for non-chemical
implementation for apple fruit may be hin-
substitutes for chemicals, such as DPA and
dered by the relatively small margin of safe
fungicides. These treatments, which may be
temperatures that do not result in damage,
applied immediately after harvest or inter-
expressed as skin browning and internal
mittently during storage, are suitable only if
breakdown (Smith and Lay-Yee, 2000).
the tissue recovers without damage after
removal from storage.
23.5 Physiological and Pathological
23.4.4.1 Low-oxygen stress Disorders
Initial low-stress treatments of 0–0.5%
23.5.1 Physiological disorders
oxygen for 5–10 days control the incidence of
superficial scald and the losses in firmness
Given the extended storage periods and the
and colour of ‘Granny Smith’ (Little et al.,
many cultivars that exist, it is not surprising
1982). Inhibition of superficial scald develop-
that a wide variety of physiological disor-
ment by initial oxygen stress has also been
ders have been identified in apple fruit.
shown for other cultivars, and it may be pos-
Susceptibility to disorders varies by cultivar,
sible to devise treatment strategies appropri-
preharvest factors and postharvest condi-
ate to the cultivar and length of storage
tions (Smock, 1977; Snowdon, 1990; Blanpied
period (Wang and Dilley, 2000). The limita-
et al., 1999; Little and Holmes, 2000).
tion to commercial adoption of initial low-
Disorders can be considered in three cate-
oxygen stress treatments may be seasonal
gories:
variations in cultivar response, and the
associated risk of fermentation. 1. Disorders that develop only on the tree.
The most important of these is water-core,
in which intercellular air spaces in the core
23.4.4.2 High-carbon-dioxide stress
and cortical tissues become filled with
Commercially, concentrations of 10–20% liquid, predominantly sorbitol (Marlow and
carbon dioxide were used to maintain flesh Loescher, 1984). Usually the occurrence of
firmness of ‘Golden Delicious’ grown in water-core is associated with advancing fruit
Washington State (Couey and Olsen, 1975), maturity and low night temperatures prior
but risk of carbon dioxide injury develop- to harvest, but a variant of the disorder can
ment limited the use of this method to other occur as a result of heat stress. In ‘Fuji’, its
regions or to other cultivars (Bramlage et al., occurrence and type of development can be
Apples - Ch.23 11/4/03 11:02 am Page 605
affected by growing region (Harker et al., (Ferguson and Watkins, 1989). In susceptible
1999). Presence of water-core in fruit at cultivars, harvest of less mature fruit can
harvest creates problems in certain cultivars, result in higher bitter pit incidence, as can
such as ‘Delicious’, because fruit with mod- excessive pruning or high temperatures
erate or severe water-core can later develop and/or water-stress conditions during the
breakdown during storage. Therefore, occur- growing season. Influences of climatic condi-
rence of water-core in ‘Delicious’ is an tions are at least partly related to their effects
indicator of the end of the harvest period on calcium concentrations in the fruit.
(Blanpied and Silsby, 1992). In contrast, Development of bitter pit during storage
water-core is desirable in the ‘Fuji’ apple results in financial loss, and a number of
because of the sweetness it imparts to the strategies have been employed to prevent
fruit, and mild or moderate water-core does its occurrence. Preharvest calcium sprays
not appear to result in the development of are commonly applied to reduce bitter pit
breakdown (Watkins et al., 1993). Grade stan- development (Ferguson and Watkins, 1989).
dards for ‘Fuji’ have recently been modified Methods to predict bitter-pit-susceptibility
so that water-core is not a grade defect for risk based on mineral concentrations (mainly
the cultivar in the USA or Canada. low calcium) at harvest (Ferguson and
2. Disorders that develop on the tree and Watkins, 1989) or infusion of magnesium
during storage. Bitter pit is a disorder char- (Burmeister and Dilley, 1994) have been
acterized by development of discrete pitting developed. Rapid cooling and CA storage
of the cortical flesh, the pits being brown and may also reduce its development during
becoming desiccated with time (Ferguson storage, but most focus has been on posthar-
and Watkins, 1989). The pits may occur vest application of calcium to fruit (Sharples
predominantly near the surface or deep in and Johnson, 1976). Calcium drenches are
the cortical tissue. An associated disorder, commonly applied to bitter-pit-susceptible
known as lenticel blotch, is also observed cultivars and in the USA are routinely
in some cultivars, such as ‘Braeburn’ and applied with DPA. Vacuum infiltration of
‘Delicious’. Sun-scald, another disorder, fruit with calcium was used commercially in
occurs in fruit on the tree, but browning/ New Zealand for several years but has now
blackening of the skin develops in storage. been phased out. Neither drenches nor vac-
3. Disorders that develop during storage. uum infiltration will totally control bitter pit
These can be divided into senescent break- in highly susceptible apples (Ferguson and
down disorders, chilling disorders and dis- Watkins, 1989; Hewett and Watkins, 1991),
orders associated with inappropriate and good preharvest management practices
atmospheres during storage. The most com- that reduce risk of subsequent disorder
mon disorders associated with temperature development should be stressed.
and atmospheres are superficial scald, soft Recommended rates for pre- and posthar-
scald, low-temperature breakdown, brown vest application of calcium vary by cultivar
core, internal browning, low-oxygen injury and region. Therefore product labels should
and high-carbon-dioxide injury. Several other be followed, in conjunction with the advice
disorders have been described (Blanpied et of the local extension specialist.
al., 1999).
Because of their commercial importance,
23.5.1.2 Senescent breakdown
factors associated with the development of
common physiological disorders are briefly Senescent-breakdown incidence is related to
summarized. the harvesting of overmature fruit and/or
fruit with low calcium concentration (Marmo
et al., 1985). It can be exacerbated by storing
23.5.1.1 Bitter pit
fruit at higher than optimal temperatures.
The incidence and severity of bitter pit are Fruit that are susceptible to breakdown
affected by cultivar, but within a cultivar bit- because of low calcium are commonly
ter pit is related to harvest date and climate drenched with calcium salts before storage
Apples - Ch.23 11/4/03 11:02 am Page 606
606 C.B. Watkins
(Sharples and Johnson, 1976), but the inci- because fruit grown in other regions may
dence of senescent breakdown can also be be susceptible to low-oxygen injury (Lau et
reduced by harvesting fruit at a less mature al., 1998).
stage, rapid cooling and reducing the dura-
tion of storage. Increased senescent-break-
23.5.1.4 Low-temperature disorders:
down incidence was shown to be related to
soft scald, low-temperature breakdown,
the climacteric in one study (Wilkinson and
brown core and internal browning
Sharples, 1967), but not in others (Blanpied,
1969; Blanpied and Silsby, 1992). However, Soft scald is characterized by irregular but
because the occurrence of breakdown is sharply defined areas of soft, light brown
highly dependent on calcium concentra- tissue, which may extend into the cortex
tions in the fruit (Marmo et al., 1985), the (Snowdon, 1990). Susceptibility of fruit to
ability to detect relationships between the soft scald is cultivar- and climate-related,
disorder and the climacteric may be con- and disorder incidence may be aggravated
founded. A fruit with high calcium concen- by harvesting overmature fruit and delays
trations may never develop senescent between harvest and cooling. Storing fruit at
breakdown, while one with low calcium 3°C rather than at lower temperatures can
concentrations may show increased disor- sometimes control the disorder (Snowdon,
der incidence with later harvest dates 1990), and DPA, used for the control of
(Watkins et al., 1989a). superficial scald, may also reduce the inci-
dence of soft scald (Wills et al., 1981). Storage
at a lower temperature following prompt
23.5.1.3 Superficial scald (syn. storage scald)
cooling can reduce the incidence of soft scald
Superficial scald is a physiological disorder on ‘Golden Delicious’ fruit.
associated with long-term storage (Wilkinson Low-temperature breakdown, brown
and Fidler, 1973; Ingle and D’Souza, 1989). It core and internal browning are affected by
was the major cause of apple-fruit loss until cultivar sensitivity to low temperatures and
the advent of postharvest treatment with the generally these disorders increase in inci-
antioxidant DPA. Cultivar, climate and dence and severity as the length of storage is
harvest date affect susceptibility of fruit to increased (Wilkinson and Fidler, 1973).
the disorder (Wilkinson and Fidler, 1973; Climate affects the sensitivity of fruit to the
Emonger et al., 1994). disorders, with more problems occurring
DPA is usually applied with a fungicide after colder, wetter growing seasons. Low-
to reduce decay incidence, and calcium salts temperature breakdown is characterized by
may also be included at the same time to markedly brown vascular bundles, brown-
reduce bitter pit or senescent breakdown. ing of flesh and a clear halo of unaffected
Both DPA use and DPA residues on imported tissue underneath the skin. In contrast to
fruit are prohibited in some countries. senescent breakdown, the affected tissues
Another antioxidant, ethoxyquin, is no longer are more likely to be firmer, more moist and
permitted for use on apples. darker in colour. Brown core (syn. core
Alternative ways of controlling superfi- flush) involves browning of the flesh, ini-
cial scald are being investigated, and stor- tially in the core area and later in the cortex,
age operators are reducing the use of DPA where it becomes difficult to distinguish
where possible. Low concentrations of oxy- from low-temperature breakdown. Internal
gen in CA storage reduce the risk of scald browning does not involve the breakdown
developing and may also permit the use of of the flesh but rather a greying of the flesh
lower DPA concentrations. Low-oxygen apparent when the apple is cut. Internal
and low-ethylene CA storage also reduce browning and core flush are often associated
scald incidence. In British Columbia, with higher concentrations of carbon diox-
Canada, 0.7% oxygen storage is used as a ide, since both can occur in CA when the
substitute for DPA treatment (Lau, 1997). carbon dioxide concentration is higher than
This technique cannot be used universally that of oxygen.
Apples - Ch.23 11/4/03 11:02 am Page 607
23.5.1.5 Low-oxygen injury to carbon dioxide injury have occurred when

DPA usage has been stopped because the
Low-oxygen injury affects fruit in a number
cultivar is not susceptible to superficial
of ways (Wilkinson and Fidler, 1973; Snow-
scald. It was not previously recognized
don, 1990; Blanpied et al., 1999). The first
that DPA reduced the incidence of both
indication of injury is loss of flavour,
external and internal carbon dioxide injuries
followed by fermentation-related odours.
(Burmeister and Dilley, 1995; Watkins et al.,
These odours may disappear if storage prob-
1997b; Fernández-Trujillo et al., 2001). These
lems are identified soon enough and severe
observations highlight the point that chemi-
injury has not occurred. Injury symptoms
cals that are technically applied for one
range from purpling or browning of the skin
reason may also have other effects that may
in red-coloured cultivars, development of
not be appreciated until the chemical is
brown soft patches resembling soft scald and
withdrawn from use. Non-chemical strategies
abnormal softening and splitting of fruit. As
to reduce carbon dioxide injury include
discussed earlier, cultivars vary greatly in
acclimatization of fruit in air before CA
response to low oxygen, and susceptibility to
storage, if this can be done without loss of
injury is influenced by a number of pre- and
quality, and initially maintaining low carbon
postharvest factors.
dioxide concentrations in the storage envi-
ronment (Wang et al., 2000).
23.5.1.6 Carbon dioxide injury
Carbon dioxide injury may be external or
internal. The external form consists of wrin- 23.5.2 Pathological disorders
kled, depressed colourless or coloured areas
restricted to the skin surface and usually on The main postharvest diseases of apples that
the greener side of the fruit. Internal forms develop in storage are blue mould, caused
are expressed as brown heart and/or cavities by Penicillium species, and grey mould,
in the flesh (Wilkinson and Fidler, 1973). caused by Botrytis cinerea. Mucor pyriformis
Occurrence of the disorders is cultivar- Fischer can cause severe losses in pears
specific, and generally external carbon diox- but is less common in apples. Botrytis and
ide injury is associated with early harvest, Penicillium species enter fruit primarily
while that of internal injury is associated through cuts, stem punctures and bruises,
with later harvest. The growing region but Penicillium expansum Link can invade
also affects susceptibility of fruit to carbon some apple cultivars via the stem during
dioxide injury (Bramlage et al., 1977; Elgar long-term CA storage (Rosenberger, 1999).
et al., 1999). Other pathogens of stored apples include the
Susceptibility to carbon dioxide injury has Colletotrichum species that cause bitter rot,
long been a problem for certain apple culti- the Botryosphaeria species that cause black
vars. It is the basis, for example, for the com- rot and white rot and Pezicula malicorticis
mercial CA recommendations for ‘McIntosh’, (Jackson) Nannf., the cause of bull’s-eye
in which carbon dioxide levels in the storage rot (Rosenberger, 1990). Decays caused by
atmosphere are kept to 2–3% for the first 4–6 Collectotrichum, Botryosphaeria and Pezicula
weeks and then allowed to increase to 5% are initiated in the field and must be con-
(Watkins et al., 2003). Several newer apple trolled by using fungicides or other disease-
cultivars, including ‘Empire’, ‘Fuji’ and management strategies during the growing
‘Braeburn’, appear to be susceptible to season.
carbon dioxide (Watkins et al., 1997b; Elgar et Blue mould and grey mould are usually
al., 1998; Volz et al., 1998), resulting in com- controlled during long-term storage by
mercial losses. Carbon dioxide injuries have postharvest application of the benzimidazole
occurred in air-stored fruit under conditions fungicide, thiabendazole (2-(4-thiazolyl)benzi-
where carbon dioxide can accumulate (e.g. midazole (TBZ) (Mertect 340F, Deccosalt 19),
warm fruit in sealed cartons). applied in combination with DPA
At least in the case of ‘Empire’, losses due (Hardenburg and Spalding, 1972). TBZ may
Apples - Ch.23 11/4/03 11:02 am Page 608
608 C.B. Watkins
be applied a second time as a line spray or in may include limited markets compared with
wax as apples are packed. Benzimidazole- field crops, liabilities associated with the value
resistant strains of P. expansum and B. cinerea of the stored crop, a high public profile for
were discovered in apple storage during apples in debates relating to food-safety issues
the mid- to late 1970s, but TBZ plus DPA and difficulties in devising shelf-stable formu-
continued to control blue and grey moulds, lations of biocontrol agents (Watkins et al., 2003).
because most benzimidazole-resistant strains Moreover, biocontrols generally cannot
of the pathogen were highly sensitive to currently provide eradicant activity against
DPA (Rosenberger and Meyer, 1985). During established infections. Using combinations of
the mid-1990s, the incidence of blue mould biocontrols and reduced rates of TBZ may be
began to increase in some apple packing- more effective than using either product alone
houses where the predominant strains of (Chand-Goyal and Spotts, 1997). When such
P. expansum had developed resistance to combinations are used, the chemical fungicide
the benzimidazole–DPA combination. Grey may provide eradicant and short-term protec-
mould is still controlled by the benzimida- tant activity necessary to prevent decay until
zole–DPA combination, presumably because the biocontrol agents become established in
this pathogen does not recycle on field bins wounds or other infection sites.
as readily as does P. expansum, and it has Regardless of the postharvest fungicide or
therefore been subjected to less selection biocontrol, good sanitation will remain
pressure for fungicide resistance (Rosen- essential for reducing inoculum on contami-
berger, 1990). Captan (N-trichloromethylthio- nated field bins and in packing-houses and
4-cyclohexene-1,2-dicarboximide; Captan 50W, storage rooms. Badly contaminated bins
Captan 80W, Captec 4L) has a postharvest should be cleaned and disinfested (steam-
registration but has proved to be only mod- cleaned) before they are reused for a new
erately effective for controlling P. expansum crop. Plastic bins may carry less inoculum
and B. cinerea. Captan residues are not than wooden bins (D.A. Rosenberger, Hudson
acceptable in some markets. Valley Laboratory, Cornell University, 2001,
Much effort has been devoted to the personal communication) and also have the
development of biocontrols for postharvest advantage of reducing bruising and abrasion
diseases of apples (Janisiewicz, 1998), but, where apples contact the sides of bins.
while many of the biocontrol agents selected Careful fruit handling, rapid cooling after
and developed to date have proved to be harvest and storage at recommended tem-
very effective in controlled tests, commercial- peratures also help to limit the development
ization of biocontrols has been slow. Reasons of postharvest decay.
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Watkins, C.B., Bowen, J.H. and Walker, V.J. (1989b) Assessment of ethylene production by apple cultivars
in relation to commercial harvest dates. New Zealand Journal of Crop and Horticultural Science 17,
327–333.
Watkins, C.B., Brookfield, P.L. and Harker, F.R. (1993) Development of maturity indices for the ‘Fuji’
apple cultivar in relation to watercore incidence. Acta Horticulturae 326, 267–275.
Watkins, C.B., Brookfield, P.L., Elgar, H.J. and McLeod, S.P. (1997a) Development of a modified atmos-
phere package for export of apple fruit. In: Proceedings of the International Congress on the Uses of
Plastics in Agriculture. Laser Pages Publishing, Jerusalem, Israel, pp. 586–592.
Watkins, C.B., Silsby, K.J. and Goffinet, M.C. (1997b) Controlled atmosphere and antioxidant effects on
external CO2 injury of ‘Empire’ apples. HortScience 32, 1242–1246.
Watkins, C.B., Nock, J.F. and Whitaker, B.D. (2000) Responses of early, mid and late season apple culti-
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phere storage conditions. Postharvest Biology and Technology 19, 17–32.
Watkins, C.B., Kupferman, E. and Rosenberger, D.A. (2003) Apple: postharvest quality maintenance
guidelines. In: Gross, K., Wang, C.Y. and Saltveit, M.E. (eds) The Commercial Storage of Fruits,
Vegetables, and Florist and Nursery Stocks. Agriculture Handbook No. 66 (revised), United States
Department of Agriculture, Washington, DC (in press).
Wilkinson, B.G. and Fidler, J.C. (1973) Injuries caused by incorrect concentrations of carbon dioxide
and/or oxygen. In: Fidler, J.C., Wilkinson, B.G., Edney, K.L. and Sharples, R.O. (eds) The Biology of
Apple and Pear Storage. Commonwealth Bureau of Horticultural and Plantation Crops, East Malling,
UK, pp. 81–87.
Wilkinson, B.G. and Sharples, R.O. (1967) The relation between time of picking and storage disorders in
Cox’s Orange Pippin apple fruits. Journal of Horticultural Science 42, 67–82.
Wills, R.B.H., Hopkirk, G. and Scott, K.J. (1981) Reduction of soft scald in apples with antioxidants.
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Handling of Fruit, Vegetables and Ornamentals. CAB International, Wallingford, UK, 262 pp.
Apples - Ch.24 11/4/03 11:02 am Page 615
24 Production and Handling Techniques for

Processing Apples
Robert M. Crassweller and George M. Greene, II

The Pennsylvania State University, Department of Horticulture, Fruit Research and
Extension Center, Pennsylvania, USA

24.2 Processed Products 616
24.2.1 Juice 616
24.2.2 Cider 618
24.2.3 Apple sauce 619
24.2.4 Slices and whole apples 620
24.3 Supply 620
24.4 Processing Industry in the Peoples’ Republic of China (PRC) 621
24.5 Important Processing Cultivars 621
24.5.1 ‘York Imperial’ 621
24.5.2 ‘Golden Delicious’ 622
24.5.3 ‘Rome Beauty’ 622
24.5.4 ‘McIntosh’ 623
24.5.5 ‘Northern Spy’ 623
24.5.6 ‘Idared’ 623
24.5.7 ‘Jonathan’ 623
24.5.8 Other cultivars 623
24.6 Fruit Attributes 624
24.6.1 Maturity 624
24.6.2 External appearance 624
24.6.3 Flesh colour 624
24.6.4 Fruit firmness 624
24.6.5 Damage and decay 625
24.7 Differences in Cultural Practices 625
24.8 Harvest and Handling Techniques 626
24.9 Delivery to the Processing Plant 627
24.10 Grade Interpretations for Payment 631
24.11 Future Directions of the Industry 631
24.1 Introduction However, in many areas more apples are

processed than are sold as fresh fruit. In
Much attention is paid to the production the USA, recent statistics from the US
practices devoted to fresh-market apples. Department of Agriculture (USDA) indicate
Apples - Ch.24 11/4/03 11:02 am Page 616
616 R.M. Crassweller and G.M. Greene, II
that, in the last 5 years, approximately 40% shipping large volumes of water. The prod-
of all the apples grown in the USA are uct is reconstituted for packaging and sale to
processed (Evans, 2000). In 1999, approxi- the consumer.
mately 2.2 million t were processed out of a Estimated 1999/2000 world production in
total crop of 5.3 million t. The era of the most selected countries is believed to be 633,000 t,
rapid growth in apple processing in the USA (70/71° Brix), 7% below the previous season’s
was the period from the end of the Second output (Rosa, 2000). The downturn mainly
World War in 1945 until the early 1970s reflected decreases in both Polish and
(O’Rourke, 1994). This time period coincided Argentine production. Production in the
with a trend where the American family USA, the world’s largest apple-juice con-
moved away from the traditional life styles centrate (AJC) producer and consumer, was
of a single-wage-earner household to a dual estimated to increase to 150,000 t.
income with less time spent on food pre- Apple juice is produced in many coun-
paration. However, in the 1970s, changes in tries. Five European countries account for
dietary patterns led to a decline in the con- trade of approximately 521,000 t year1
sumption of processed fruits and vegetables. (Neubert and Lee, 1999). This quantity
In general, the processing industry has been includes both production and refiltration of
geographically tied to locations where there juice initially produced in other countries.
were ample local supplies of apples. These countries are mainly exporters, con-
suming only 35% of the traded product and
exporting the remainder. Poland, Italy and
24.2 Processed Products Germany were the top three European AJC-
exporting countries in 1998, according to the
24.2.1 Juice Food and Agriculture Organization (FAO)
(Table 24.1). Germany is a major producer
Apple-processing products are varied and of AJC, producing 73,872 t in 1997/98
depend upon the state or country and the (Luetzenkirchen and Frimmersdorf, 1998). In
economic status of the area. The primary Poland, approximately 60% of the apples
processing product produced in the world is produced are processed. Processed products
apple juice. Apple juice is a clarified product include AJC, fruit beverage, wine and jam.
that is pasteurized to make it shelf-stable. In Germany is the leading destination for
international trade, apple juice is usually Polish AJC, accounting for more than 80% of
concentrated to 70° Brix or higher to reduce the total AJC exports. The German AJC
Table 24.1. Top ten concentrated-apple-juice exporting and importing countries in 1998 (t) (from the
Food and Agriculture Organization of the United Nations available at http://apps.fao.org/). Production
from the Peoples’ Republic of China is not included in this summary.
Exports Imports
Rank Country Volume Rank Country Volume
1 Poland 104,497 1 USA 219,971

2 Italy 79,392 2 Germany 123,665
3 Germany 73,397 3 UK 35,913
4 Austria 58,253 4 Italy 33,227
5 Argentina 54,732 5 Austria 22,566
6 Turkey 51,275 6 The Netherlands 16,203
7 Chile 33,756 7 Canada 14,856
8 USA 25,005 8 Belgium–Luxemburg 10,754
9 Moldova, Republic of 18,630 9 Denmark 6,640
10 Switzerland 9,060 10 France 6,295
Total all Total all

FAO member nations 551,853 FAO member nations 515,203
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11/4/03
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Processing Apples: Production and Handling Techniques
Page 617
Table 24.2. Five-year US fresh and processed apple production (’000 t) (from Fruit and Tree Nuts, FTS-290, National Agricultural Statistics Service, USDA).
Total Fresh Total Juice Per cent

Year production market processed and cider Tinned Frozen Dried Other processed
1995 4519.3 2649.1 1870.2 1150.3 586.0 138.3 151.4 35.4 41.4
1996 4829.3 2815.4 2013.9 990.3 587.0 121.5 143.6 27.8 41.7
1997 4610.6 2637.4 1973.1 973.0 679.9 158.3 121.1 81.6 42.8
1998 4916.9 2908.7 2008.3 1127.3 532.4 120.7 149.6 43.1 40.8
1999 4713.6 2705.3 2008.3 1115.8 611.6 97.7 126.2 57.1 42.6
617
Apples - Ch.24 11/4/03 11:02 am Page 618
industry also depends on supplies from Italy. Outside the USA, ‘cider’ is the fermented
Other significant sources include Turkey, the juice of the apple and has been known for
Czech Republic and Moldova. The USA centuries. Its production and use probably
remains the largest single export market for originated in the Trans-Caucus region. Cider
German AJC, while the USA and Germany spread through the Greek and Roman
were the top importing countries. Most com- worlds, but the product was not as popular
mercial apple cultivars will produce an as wine. Eventually it became associated
acceptable juice if they are blended. The qual- with the more northern regions of Europe,
ity and taste of the juice, however, is directly where grapes were not as successful. The
related to the cultivar and maturity of the major cider-making areas now are England,
fruit. Hazy or cloudy AJC is often the result north-western France, Spain, Germany and
of high levels of starch in the fruit from being Switzerland. The industry has grown in
harvested too early. Juice from less devel- these areas partly as a tourist attraction, with
oped countries may be cloudy due to their ‘Routes des cidres’ being developed simi-
inability to provide adequate filtration larly to the wine industry (Rowles, 2000).
and/or heating (Neubert and Lee, 1999). The major production areas in the UK are
In the USA approximately 54% of the Hereford, Somerset, Devon, Avon, Norfolk
processed fruit is made into juice (Table 24.2), and Sussex. Recently in the USA, there has
followed by tinned, frozen, dried and other been an increase in cider production, with
uses. On a local basis, in nearly every state of the product being marketed to a specific
the USA, small apples and those not meet- age-group (Rowles, 2000).
ing fresh-market standards are processed Traditionally, apples grown for cider were
into fresh non-pasteurized juice, which is produced on standard trees in small
typically referred to as sweet or fresh cider. orchards with the grass under the trees being
This cider is sold locally by the farmer or to grazed by livestock as part of a mixed self-
local stores. It must be refrigerated and has a sufficient farm (Williams, 1992). There was
shelf-life of no more than 2–3 weeks. minimal management or pest control and
Recently, regulations have been proposed fruit may have been harvested from the tree
by the Food and Drug Administration of the or gathered from the ground. Traditional
USA that will mandate pasteurization for all orchards planted on seedling rootstocks have
juice producers that sell cider to wholesale slowly been replaced with more intensively
markets. Under the regulations, juice proces- managed semi-intensive orchards in recent
sors must develop and implement a hazard years. Recent production figures compiled
analysis and critical control point (HACCP) by the Ministry of Agriculture, Forestry and
plan that includes control measures suffi- Fisheries in the UK indicate that average pro-
cient to achieve a 5-log (99.999%) reduction duction for 1996–1999 was around 75,000 t
in harmful pathogens. Producers that sell and that the average price was around
directly to the consumer will not have to US$141 t1 (Lawton, 2000) on an estimated
pasteurize but will have to place a warning area of 5043 ha.
label on their containers indicating that the Cider apples are distinct cultivars and are
product was not pasteurized. chosen for their fruit qualities (Table 24.3).
The cultivars can be classified into four
groups: (i) sweet – low both in tannin (0.2%)
24.2.2 Cider and in acidity (< 0.45%); (ii) bitter-sweet –
high in tannin (> 0.2%) but low in acidity;
Sweet cider is not to be confused with the (iii) bitter-sharp – high in both tannin and
fermented cider that is produced predomi- acidity; and (iv) sharp – low in tannin, high
nantly outside the USA. Fermented cider is a in acidity. Tannin level is often regarded as
shelf-stable product that has been fermented the distinguishing feature of cider apples. It
to increase the alcohol content and is typi- gives the fresh fruit a bitter and astringent
cally sold in restaurants or pubs. In the USA taste. Tannins also help prevent breakdown
it is generally known as hard cider. of the apple pulp during processing and
Apples - Ch.24 11/4/03 11:02 am Page 619
Processing Apples: Production and Handling Techniques 619
Table 24.3. Cider-apple cultivars grown in the UK by taste group.
Bitter-sweet Sharp
‘Ashton Bitter’ ‘Backwell Red’
‘Ashton Brown Jersey’ ‘Bramley’s Seedling’
‘Ball’s Bittersweet’ ‘Brown’s Apple’
‘Belle Fille De La Manche’ ‘Cider Lady’s Finger’
‘Black Dabinett’ ‘Crimson King’
‘Brimley Bittersweet’ ‘Frederick’
‘Brown Snout’ ‘Improved Lambrook Pippin’
‘Brown Thorn’ ‘Reinette O’bry’
‘Bulmers’ Norman’ ‘Severn Bank’
‘Burrowhill Early’ ‘Stembridge Cluster’
‘Chisel Jersey’ ‘Tom Putt’
‘Coat Jersey’
‘Collington Big Bitters’
‘Dabinett’
‘Doux Normandie’
‘Dove’ Sweet
‘Dymock Red’ ‘Court Royal’
‘Ellis Bitter’ ‘Dunkerton Late Sweet’
‘FillBarrel’ ‘La Bret’
‘Harry Masters Jersey’ ‘Morgan Sweet’
‘Improved Dove’ ‘Northwood’
‘Major’ ‘Sweet Alford’
‘Maundy’ ‘Sweet Coppin’
‘Medalle D’or’ ‘Tall Sweet’
‘Michelin’ ‘Taylors’
‘Muscadet De Dieppe’
‘Nehou’
‘Omont’
‘Osier’
‘Perthyre’ Bitter-sharp
‘Reine De Hatives’ ‘Breakwell’s Seedling’
‘Reine des Pommes’ ‘Broxwood Foxwhelp’
‘Rougette Douce’ ‘Foxwelp’
‘Somerset Redstreak’ ‘Genet Moyle’
‘Stable Jersey’ ‘Improved Redsteak’
‘Stembridge Jersey’ ‘Kingston Black’
‘Tardive Forestiere’ ‘Porter’s Perfection’
‘Tremlett’s Bitter’ ‘Red Foxwhelp’
‘Vagon Archer’ ‘Stoke Red’
‘Vileberie’
‘White Jersey’
‘White Norman’
‘Yarlington Mill’
makes it easier to press cider cultivars than product. Apple sauce is produced by peeling
fresh-market cultivars. and coring the fruit and then slicing them
into small irregular pieces, to which sugar
or maize syrup may or may not be added.
24.2.3 Apple sauce The mixture is then precooked, usually by
passing through a pressurized steam tunnel
Apple sauce is a uniquely North American for 4–5 min until the mixture temperature
product. A mixture of cultivars is usually reaches about 96°C. Once the fruit is cooked,
used to achieve a consistent and uniform it is passed through a finishing machine that
Apples - Ch.24 11/4/03 11:02 am Page 620
regulates the sauce grain and removes any 24.3 Supply

large coarse materials. If the sauce is placed
directly into tins and the mixture tempera- In the eastern USA (primarily Pennsylvania,
ture is maintained at 93°C, no further cook- Virginia, West Virginia, New York and
ing is needed. If the temperature of the Michigan), apples grown for processing are
mixture drops, however, then another cook- utilized for apple slices, apple sauce, whole
ing step is needed to kill any pathogens that baked apples, spiced apple rings and apple
may be present. juice. The processing industry developed in
Processors look for specific characteristics these states as the market for processed
in apples, such as Brix (11–24°), sugar : acid products expanded and as growers increased
ratios (25–60), aroma, yellow or white flesh, production (Plate 24.1).
variable grain or texture and good water- In the western USA (primarily Washing-
holding capacity. Cultivars with high water- ton, Oregon and California), apples are
holding capacity allow the addition of water diverted into processing uses as a result of
to increase the yield of the end-product. The fruit not meeting fresh-market quality stan-
US standards for grade of tinned apple sauce dards. About 20–30% of the apples end up
are based upon five attributes: colour, con- at processing plants (Hansen, 1997). Few
sistency, absence of defects, flavour and fin- growers in the Pacific north-west region are
ish. Finish is primarily due to the size and growing specifically for processing. While
distribution of the apple tissue in the sauce, the majority of fruit ends up in juice, some
which is cultivar-dependent (Nogueira et al., higher-quality fruit may be utilized for other
1985). products. Typical cultivars utilized include
‘Golden Delicious’, ‘Granny Smith’, ‘Rome
Beauty’, ‘Winesap’, ‘Gala’ and ‘Fuji’.
Processors in Washington look for fruit that
24.2.4 Slices and whole apples
is a minimum of 64 mm in diameter.
Premiums are sometimes paid for internal-
Another product is the production of whole
quality parameters, such as high acid. Prices
baked and sliced apples. These are often
fluctuate depending upon supply and
processed in tins and utilized in pie fillings.
demand. In the eastern USA, prices are
Slice packs generally consist of a single culti-
usually set at the beginning of the season. An
var and are therefore not blended. Trim
economic disadvantage for Pacific north-
waste must be minimal and firmness and
west growers is the added cost for storage.
texture are important qualities for cultivars
In Canada, Nova Scotia’s processing
destined for this product. Uniformity of size,
industry is closely aligned with a specific
shape and core diameter are important to
bakery company that prefers ‘Northern Spy’
obtain maximum yield when fruit are apples. The remainder of the processed pro-
mechanically peeled and cored. The apple duction consists of packing-house fruit that
slices can also be further processed by de- did not meet fresh standards and a small
hydrating and freezing. In recent years, hard-cider industry in eastern Canada. In
small local processing operations have been British Columbia, a single company domi-
established to supply nearby bakeries for nates the processing market and products
pastry uses. Apples destined for sauces and are primarily made from fruit that did not
slices receive a higher premium than those meet fresh-market standards (Advanced
processed into juice. Resource Consulting Ltd, 1999).
Other products are made depending While the tonnage of processed apples is
upon the individual processor’s ability to high, the value of the crop is not necessarily
diversify his/her product line. Apple butter, on a par with fresh-fruit uses. In 1998, the
spiced apple rings and apple jelly are consid- estimated value of apples was just over
ered speciality products made from apples US$1.3 billion, of which US$1.1 billion was
and constitute less than 1% of total products for fresh fruit and only US$0.2 billion was
produced (Root, 1996). attributed to processed fruit (Evans, 2000).
Apples - Ch.24 11/4/03 11:02 am Page 621
24.4 Processing Industry in the People’s and, in recent years, ‘Fuji’. The New York
Republic of China (PRC) processing industry is based upon
‘McIntosh’, ‘Rhode Island Greening’, ‘Wayne’
In recent years, apple production in the PRC and ‘Northern Spy’ (S. Hoying, New York,
has dramatically increased. Acreage in the USA, 2000, personal communication).
country is estimated to be around 2.44 The Michigan processing industry breaks
million ha (Rutledge, 2000). It is estimated their processing cultivars down into three
that only about 5–10% of each year’s apple categories (D. Ricks, Michigan, USA, 2000,
crop is processed, the dominant product personal communication). At the extremes
being juice or juice concentrate. While ‘Fuji’ are those grown only for processing and
is the most important apple cultivar grown those that are grown only for fresh-market
in the PRC, the cultivar ‘Qinguan’ is the consumption. In between are the dual-
most popular among the country’s AJC purpose cultivars that can be grown for
producers. The rise in apple-juice production either the fresh or the processing market. The
is largely a result of foreign joint-venture cultivars grown primarily for processing
investments with companies such as Dole/ include ‘Northern Spy’, ‘Winesap’, ‘Rhode
Tropicana, Great Lakes, Kirin and Ronghzi. Island Greening’ and ‘Mutsu’. The-dual
These companies buy local deciduous fruit purpose cultivars are ‘Jonathan’, ‘Golden
to manufacture into juice for the domestic Delicious’, ‘Rome Beauty’, ‘Idared’ and
market. Total AJC produced in 1998 was ‘Jonagold’. Processed products from these
90,900 t and utilized 900,000 t of apples (4.6% two categories include apple sauce, tinned
of the total crop). In 1999, this increased to apple slices, frozen slices and pie filling. The
153,059 t. The majority of the juice that was fresh-market cultivars can also be diverted
exported went to Japan and Europe. into processing, but, as in the western USA,
It is believed that there are approximately these are primarily packing-house culls that
55 processing plants in the country, with the are made into juice. Culls are fruit that are
largest number in the Shandong (22 plants) unmarketable due to small size or quality
and Shaanxi (17 plants) provinces. Unlike the defects, such as poor colour and disease and
industry in the USA, the juice production in insect damage.
the PRC is seasonal, starting in August and In eastern European countries, such as
running until about January or February, Poland, older cultivars, such as ‘Antonovka’,
when the supply runs out. Apples are not ‘McIntosh’ and cultivars of local importance,
stored for later use. The amount of juice pro- dominate the processing market. Other
duced is expected to continue to increase for cultivars recommended in Poland include
the short term, as only 60–70% of the PRC’s ‘Landsberge’, ‘Wealthy’, ‘Golden Delicious’,
apple trees are bearing. ‘Idared’ and ‘Warta’ (A. Czynczyk,
Skierniwice, Poland, 2000, personal commu-
nication). Most of the dominant processing
24.5 Important Processing Cultivars cultivars developed as chance seedlings. In
the USA many of the prominent processing
All apple cultivars grown can be used to cultivars are also among the leading culti-
some extent for processing. However, there vars in production (Table 24.4).
are certain characteristics that make some
cultivars more desirable for different end-
products. Cultivars used for processing vary 24.5.1 ’York Imperial’
by region. The large mid-Atlantic region,
which encompasses the states of ’York Imperial’ was a chance seedling found
Pennsylvania, West Virginia, Maryland and in York, Pennsylvania. It was discovered
Virginia, primarily utilizes ‘York Imperial’, when schoolboys passing by a seedling tree
‘Golden Delicious’ and ‘Rome Beauty’ for in late March picked up the dropped fruit
processing. Other cultivars that processors and found how firm the fruit remained
utilize include ‘Granny Smith’, ‘Winesap’ (Rollins, 1989). It is a late-maturing cultivar,
Apples - Ch.24 11/4/03 11:02 am Page 622
Table 24.4. US apple production by cultivar (’000 t) (from US Apple Association).
2000
Cultivar 1995 1996 1997 1998 1999 forecast
‘Delicious’ 1910 1982 1720 2084 1575 1701

‘Golden Delicious’ 673 680 653 757 625 674
‘Fuji’a 197 248 300 394 312 388
‘Granny Smith’a 272 312 310 359 370 374
‘Gala’a 125 159 181 233 275 308
‘Rome Beauty’ 334 257 276 265 272 250
‘McIntosh’ 248 212 259 198 281 214
‘Jonathan’ 142 87 104 114 126 101
‘York Imperial’ 106 78 87 71 98 88
‘Idared’ 107 75 96 98 113 88
‘Empire’ 83 83 95 82 102 82
‘Cortland’ 50 47 49 45 56 44
‘Rhode Island Greening’ 62 46 43 42 49 40
‘Newtown’ 65 58 56 54 32 35
‘Stayman’ 41 27 31 30 34 30
‘Northern Spy’ 43 25 35 35 50 29
‘Winesap’ 33 14 18 18 19 13
‘Gravenstein’ 18 9 9 8 4 4
All others 291 312 361 397 408 381
Totalb 4801 4709 4683 5283 4800 4843

a Includes only western production. Eastern and mid-west production is included in ‘all others’.
b Sum of cultivars may not add to total due to rounding of individual cultivars.
being harvested approximately 170–180 days Nursery. ‘Golden Delicious’ fruit are
after full bloom. The fruit are characterized regarded as being sweet and semi-firm and
as being firm to hard, with a creamy-yellow they store well and are non-browning when
flesh. The firmness and colour of the fruit cut. The fruit is harvested 135–150 days
provide superior processed sliced products after full bloom. Horticulturally the tree is
that hold their shape. ‘York Imperial’ has a classified as an easy tree to prune and train.
long storage life and is resistant to bruising. It is rated as very good to excellent for most
The fruit has a high water-holding capacity, processed products (Way and McLellan,
which results in a high processing yield for 1989). One of its major advantages is its
sauce. Disadvantages of the cultivar include versatility in making processed products and
its lopsided shape, which can affect the peel- its ability to be blended with other cultivars
ing process, and a lack of aroma. Culturally, to improve the uniformity of the finished
the tree is vigorous, with a tendency to product.
biennial bearing and a propensity to develop
calcium deficiency-related disorders, such as
cork spot and bitter pit. 24.5.3 ‘Rome Beauty’
‘Rome Beauty’ was discovered as a sucker

24.5.2 ’Golden Delicious’ growing from below a graft union in the
early 1800s in Ohio in the USA. It has
’Golden Delicious’ was a chance seedling become the fifth most popular cultivar in the
found in West Virginia in the USA on the USA. Its major advantage is good adaptation
Andrew Mullins farm (Maas, 1970). It is not to a wide range of growing conditions,
related to the ‘Delicious’ cultivar, other than heavy annual bearing, large fruit size and
both being commercialized by Stark Brothers late bloom. When young, the tree has an
Apples - Ch.24 11/4/03 11:02 am Page 623
upright growing habit, which gradually tages of the cultivar are the large fruit size,
develops into an acrotonic growth habit as it yellow flesh colour, flavour and late-bloom-
matures. The fruit can be used either fresh or ing tendency. Disadvantages of ‘Northern
for processing, although neither is of high Spy’ are its extreme lack of precocity and its
quality. It is less desirable for apple sauce vigorous growth and biennial-bearing ten-
than most cultivars, because of poor flesh dency. Use of dwarfing rootstocks is manda-
colour (Way and McLellan, 1989). Fruit tory to reduce the growth and lack of early
matures approximately 165–170 days after production.
full bloom but can hang longer on the tree.
Storage life is very good. Its best processing
use is in the making of whole baked-apple 24.5.6 ’Idared’
products. ‘Lawspur’ and other high-colour-
ing red strains of ‘Rome Beauty’ are some- ‘Idared’ was produced by a cross made in
what less desirable for use in processing, Idaho of ‘Jonathan’ ‘Wagener’ in 1942. It is
because of the tendency for the development a dual-purpose fruit, being grown for both
of red coloration in the flesh of the fruit. the fresh and the processed markets. The
bright, shiny, red fruit ripen 145–160 days
after full bloom. The major advantages of
24.5.4 ’McIntosh’ this cultivar are its large round fruit, pre-
cocity and long storage life. Disadvantages
‘McIntosh’ was discovered by John McIntosh include the tree susceptibility to both fire
in about 1811 in Dundela, Ontario, in blight (Erwinia amylovora (Burr.) Winslow et
Canada, but it was not until about 1900 that al.) and powdery mildew (Podosphaera leu-
the cultivar became widely known (Upshall, cotricha (Ell. & Ev.) E.S. Salmon).
1970). ‘McIntosh’ fruit ripen 120–135 days
after full bloom and the cultivar is con-
sidered an annual bearer. It is known to be 24.5.7 ’Jonathan’
a cold-hardy cultivar, having survived cold
winters when ‘Baldwin’ trees were killed. ‘Jonathan’ is believed to be a chance seedling
Today ‘McIntosh’ is the seventh most widely of ‘Esopus Spitzenberg’ found in Kingston,
grown cultivar in the USA. It is also popular New York, around 1825 (Larsen, 1970). It is
in eastern European countries, such as well adapted to the eastern and midwestern
Poland. The flesh of ‘McIntosh’ is distinc- regions of the USA as well as central Europe,
tively white and produces a whitish-coloured although its production has declined com-
processed product. ‘McIntosh’ is in many pared with previous years. It is, however, still
cases grown as a fresh fruit cultivar and fruit widely grown for processing in Michigan.
that do not meet fresh-market standards are Apple sauce made with ‘Jonathan’ has a
diverted into processing. good texture and an appealing yellow colour.
Small fruit size is a drawback, resulting in
less efficiency for mechanical peelers and
24.5.5 ’Northern Spy’ higher trim and core waste. The tree is very
susceptible to fire blight, powdery mildew
‘Northern Spy’ was a chance seedling found and cedar apple rust (Gymnosporangium
in approximately 1800 in western New York juniperi-virginianae L.).
in the USA, but it was not until about 1840
that it was recognized outside its small area
of origin. Fruit from ‘Northern Spy’ ripen 24.5.8 Other cultivars
approximately 150–160 days after full bloom.
The fruit are large and distinctly ribbed. The Cultivars of lesser importance include
best processed products produced from the ‘Gravenstein’, grown primarily in California
fruit include slices, either tinned or frozen, and producing an excellent product for
and whole baked apples. The major advan- apple sauce and pie filling. ‘Rhode Island
Apples - Ch.24 11/4/03 11:02 am Page 624
Greening’ is an old cultivar that is still 76 mm apple has 60% of the fruit that can be
produced in New York and Michigan but is converted into slices (Cooper, 1988).
losing favour. It is commonly known as
‘Greening’ and, as the name implies, is green
in colour when harvested for processing. 24.6.1 Maturity
Apple sauce made with this cultivar has a
high yield due to its high water-holding Soft fruit do not withstand the rigours of
capacity. ‘Granny Smith’ is a chance seedling mechanical peeling, cooking and thermal
from Australia that in recent years has been processing as well as firmer fruit. Overripe
increasingly used for processed products. fruit also do not have as high a juice yield
The late-harvested fruit make excellent pie as firm mature fruit. Less mature fruit will
fillings. ‘Twenty-Ounce’ is a large-fruited produce a grainy sauce, which is a positive
cultivar that is produced in New York but aspect (Lanza and Kramer, 1967).
with declining production. Way and McLellan
(1989) indicate that it can be used without
blending for sauce and that it has a high 24.6.2 External appearance
yield due to its large fruit size. Other culti-
vars that are used for processing include Generally speaking, apples destined for pro-
‘Mutsu’, ‘Jonagold’, ‘Monroe’ and ‘Stayman’. cessing do not need to meet the same
With the exception of ‘Monroe’, processing is appearance standards as those established
usually a secondary market, as these culti- for fresh-market fruit. Colour, which is an
vars all have a higher fresh-market value. important grade factor for fresh fruit, is less
important. Red skin, however, can limit use
in processing when the skin colour ‘bleeds’
24.6 Fruit Attributes into the flesh of the fruit. This is particularly
true with certain strains of the cultivar
The quality of the fruit delivered to the ‘Rome Beauty’. Further complicating the
processor ultimately affects the quality of the process is the fact that red anthocyanin pig-
end-product. LaBelle (1981) has described ments are unstable and turn brown during
the important quality characteristics of the long-term storage of tinned products, ren-
raw product that are desired by the pro- dering them unusable for producing a pink
cessors. Factors that have an impact on the sauce or juice (LaBelle, 1981).
end-product include ripeness, bruising,
decay, soluble solids, flesh colour, total
solids, total acid, pH, organic flavour 24.6.3 Flesh colour
compounds, tannins and juiciness. Specific
attributes may be more or less important, The flesh colour of apples varies by cultivar
depending upon the intended product. For and age of fruit. The flesh of immature fruit
baked apples, firmness, damage and core may have a distinct green tinge, while over-
size are important, while, for sliced apples, mature fruit may be deep yellow. The indi-
damage and core size are less important. vidual preference of the processor for the
Fruit size and core size have an impact on end-product colour can dictate the cultivar
the product yield and hence the profitability preference for that processor. Certain com-
of the processor. Therefore, the larger the panies produce a more yellow sauce, while
fruit diameter, the fewer units of fruit that others prefer a whiter or cream-coloured
have to be peeled to produce a given amount sauce.
of useful flesh. It takes 45 64 mm (2 in.)
apples to produce 10 lb of slices, while it
takes only 26 76 mm (3 in.) apples to make 24.6.4 Fruit firmness
4.5 kg (10 lb.) of slices. Therefore, an apple
that is approximately 64 mm has only 51.3% Fruit firmness is one of the major criteria uti-
of its flesh that is used to make slices, while a lized by classifiers at the receiving station to
Apples - Ch.24 11/4/03 11:02 am Page 625
determine the immediate placement of the and defects. Light exposure is still important
fruit. The firmest fruit are usually directed to ensure adequate flower formation and
towards long-term controlled-atmosphere return bloom, and the trees therefore need to
(CA) storage, while the softest fruit can be have a fairly open structure. However, in
directed into immediate use in the process- many instances the pruning is not as detailed
ing plant. Fruit firmness also influences the as is demanded by fresh-market standards,
final product that is to be made from the since fruit colour at harvest is not a criterion
fruit. Apples destined to be processed into for payment. Good processing growers try to
tinned slices or pie filling must be firmer perform some pruning in all their blocks each
than those to be used for sauce or juice. year, but may only do a more detailed pruning
Minimum firmness for slices is 53 N (12 lb.) – every 2–3 years. In older orchard blocks on
less than this and the slices will not retain seedling rootstocks, it was common to estab-
their integrity after cooking. lish tree height by using tree-topping equip-
ment. This may or may not be followed up in
the same year with a few detailed cuts. Harper
24.6.5 Damage and decay (2000) estimated costs for pruning large, stan-
dard, processing trees on a per-tree basis to be
Fruit that has an excessive amount of bruis- US$4.00–5.00, due primarily to the large tree
ing, hail damage, corking or insect feeding is size.
downgraded during inspection. Most The conversion to high-density plantings
processors list bruising as the biggest single on dwarfing rootstocks for processing
problem. Apart from direct fruit damage orchards has not been as rapid as the conver-
from insect feeding, there may also be an sion in fresh-market orchards. The slow con-
impact upon processor returns. Hull and version is probably due to three factors. First,
Rajotte (1988) showed that fruit placed in many of the established processing orchards
storage with damage from insect feeding are old and were established many years
tended to have an increased susceptibility to before a great variety of dwarfing rootstocks
fungal disorders and to a shortened storage were available. It is not unusual to have pro-
life. The damaged fruit were still classified cessing orchards that are in excess of 35 years
as US No. 1 and therefore did not have an old. Secondly, since external fruit appearance
impact on the fruit grower’s financial return is not as critical, the need for small, compact,
but would have had an effect upon the tree canopies that produce high-coloured
flexibility of use and yield for the processor. fruit is not as great as in fresh-market pro-
duction systems. In a 10-year study of ‘York
Imperial’ on seven different rootstocks,
24.7 Differences in Cultural Practices Greene et al. (1997) found that total cumula-
tive yield for the years 1991–1996 (fifth to
Cultural practices for growing processing tenth year) varied from 85.6 to 125.3 t ha1.
apples are not very different from practices Early in the life of the orchard, it appeared
utilized for fresh-market apples. In the case that some of the smaller trees had higher
of fruit that is directed to the processor as a yield efficiencies than some of the larger
salvage from fresh fruit, there would be no trees. Thirdly, there has not been the need to
differences in their cultural practices since change to new, higher-value, apple cultivars.
the fruit utilized is the result of off-grade or An advantage that the processing grower has
poorer quality fruit that was initially is the stability of the cultivars. Much of
intended for the fresh market. today’s change in apple systems is predicated
Fruit that is grown intentionally for the on the need to plant new cultivars that bring
processor does have a few changes in cultural a high price in the market, hence the need to
practices, but not as many changes as one continually replant to stay current with mar-
would expect. As mentioned previously, the ket pressures. Processing orchards do not
processors pay a premium for larger-sized need to replace cultivars and therefore can
fruit that have a minimal amount of bruising have a longer lifespan.
Apples - Ch.24 11/4/03 11:02 am Page 626
The number of pesticide applications is fruit strictly for processing are estimated to
only slightly lower for processed fruit, but be approximately US$740 ha1 cheaper than
there is less concern about fruit finish and the costs of production for fresh-market
the materials utilized may be less expensive. apples (Table 24.5). The budget was based
Economic injury thresholds are higher. upon a mature orchard at a density of 190
Fertilization practices may be different in trees ha1 (6.0 m 8.5 m) as compared with
orchards where the fruit is grown for pro- a fresh market orchard at 672 trees ha1
cessing. Since payment is based upon fruit (3.0 m 4.9 m). These costs are not based on
size and total weight, the fruit grower may a specific orchard but are meant to be gener-
frequently apply a higher rate of nitrogen to alized estimates based on grower and
the trees in an attempt to increase fruit size. university personnel input. Potential returns
Increasing nitrogen does have a detrimental are lower due to the lower prices paid for
affect upon fruit firmness, colour and stora- processed fruit. Prices quoted by processors
bility, but these factors may not have an (Table 24.6) can vary, depending upon
impact on the overall grower return. supply and demand and the world price for
The major difference in cultural practices apple concentrate. Harvest costs are esti-
between fruit grown for processing and that mated at US$1.35 per bushel (19 kg unit).
grown for the fresh market is in the use of Fruit grown specifically for processing do
special growth-regulator sprays. Where not require the added cost of grading and
fresh-market fruit may be treated with gib- packing that is required for fresh-market
berellins to increase fruit size and fruit fruit (Crassweller, 1995).
length-to-diameter ratio, it would not be
economically justified to do the same on
processing cultivars. Nevertheless, many fruit 24.8 Harvest and Handling Techniques
that are diverted to the processor due to in-
ferior quality may have been treated if the Processing companies require a consistent
remainder of the crop had been destined for supply of good-quality fruit. Fruit must be
the fresh market. free of insect larvae. At inspection, there is
Production costs for orchards that grow zero tolerance for insects and any amount
Table 24.5. Mature processing apple orchard budget, 77 trees acre1 (190 trees ha1), Pennsylvania,
1998. Estimated operation and input costs per acre (US$).
Month Tractors and Materials or Total

Operation performed equipment Labour service costs
Pruning and training February – 308.00 – 308.00

Liming February – – 12.50 12.50
Applying herbicides March 1.72 3.12 12.63 17.46
Fertilizing May 2.10 1.00 23.90 27.00
Bee rental May – – 25.00 25.00
Chemical thinning May 1.87 1.98 17.25 21.10
Mowing Season ( 5) 22.65 18.95 - 41.60
Applying pesticides Season ( 12) 22.44 23.76 456.18 502.38
Mouse control November 2.10 1.00 10.00 13.10
Totals 52.88 357.81 557.46 968.15
Interest on operating
capital – – – – 46.25
Land charge – – – – 150.00
Total specified costs 1164.40

Apples - Ch.24 11/4/03 11:02 am Page 627
Table 24.6. Returns above specified growing and harvesting costs under various price and yield
combinations (US$).
Orchard Yield
block
pricea 200 400 600 800 1000 1200
(US$ bu1) bu acre1 bu acre1 bu acre1 bu acre1 bu acre1 bu acre1
1.50 (1114) (1064) (1014) (964) (914) (864)

2.00 (1014) (864) (714) (564) (414) (264)
2.50 (914) (664) (414) (164) 86 336
3.00 (814) (464) (114) 236 586 936
3.50 (714) (264) 186 636 1086 1536
4.00 (614) (64) 486 1036 1586 2136
4.50 (514) 136 786 1436 2086 2736
5.00 (414) 336 1086 1836 2586 3336
5.50 (314) 536 1386 2236 3086 3936
Values in parentheses indicate returns below production costs.

a Picked in bins in orchard. Average price received for all apples (US$).
bu, bushel.
found by the inspector will automatically the sod, as well as choosing slow-growing
cause the entire load to be rejected. The grass species.
biggest single problem processing compa-
nies have is bruised fruit (Cooper, 1988).
Most processing companies now operate 12 24.9 Delivery to the Processing Plant
months a year and therefore utilize CA stor-
age. Bruised apples have a shorter storage Figure 24.1 is a general flow diagram of how
life and companies may, therefore, pay a pre- a processing plant might receive, categorize
mium for fruit with lower amounts of bruis- and inspect and grade apples. At the pro-
ing. Growers are continually cautioned that cessing plant, the truck is first driven to a
they should not have two different picking check-in station and weighed (Fig. 24.2). The
standards for fresh and processing fruit. The next stop is a subjective sampling, where a
USDA grade allows for no more than 10% quality-control person collects small sub-
bruising. Due to the need for minimal samples to measure fruit maturity and to
bruised fruit, machine harvesting of fruit has check for major quality defects. This is done
not been developed. There is interest in to determine how long the fruit can be stored
utilizing machines if one can be developed (Fig. 24.3). The maturity of the fruit is
that can minimize bruising. Currently in the typically evaluated based on fruit firmness,
USA, most fruit are harvested by hand and starch rating, visual inspection and, in some
placed into large wooden bulk bins, which cases, internal ethylene. Based upon the
typically hold 20–25 bushels (381–476 kg). results of these tests, the fruit may then be
The fruit are then loaded by fork-lift on to designated for immediate processing (most
flat-bed trucks for transportation directly to mature), regular cold storage or long-term
the processing receiving station. Williams and CA storage. Each bin is labelled with a bar-
Copas (1992), however, indicate that there code, which contains information about the
are several types of mechanical harvesters in grower, results from the maturity tests and
England for cider-apple production. These the cultivar name (Fig. 24.4).
include ‘shake-and-catch’ machines and The next step is the inspection to determine
sweepers that remove fruit from the orchard. the grade and size distribution of the entire
When using mechanical harvesting equip- lot, which will be used for payment. In the
ment, it becomes more important that the USA, grades utilized can be either USDA
orchard floor is well managed. Management grades or other grades agreed to by the
includes the proper levelling and mowing of processor and grower. A random sample of
Apples - Ch.24 11/4/03 11:02 am Page 628
Payment
determined
Sample Graded
bins and sized
Immediate
use
Truck Subjective Grade Remainder Regular

weighed inspection inspection of load storage
CA
storage
Test
results
Solid lines indicate movement of fruit

Dashed lines indicate movement of information
Fig. 24.1. Flow diagram of a typical fruit-receiving process at a processing plant.
Fig. 24.2. The trucks with loaded bins are driven over a scale to determine the weight of the fruit being
delivered and then the truck is weighed after the bins are unloaded.
bins is chosen, based upon a random-number fruit from the bins. In some instances, the
generator and the total number of bins on the entire bin is dumped over a sizer with a gap
truck. These bins are removed for grading. cut to drop out randomly a sample of fruit. In
There is no one established method for collect- another system, the bin is covered with a cor-
ing a sample from the bins, due to the differ- ner that has a square 30 cm 30 cm hole.
ent nature of the processing plants. However, When the bin is inverted, the apples from the
it is the responsibility of the person or agency uncovered area drop out to be graded. The
assigned to conduct the inspections to collect a location of the uncovered square is varied
good representative sample of the lot. with each bin. Some processing companies
Bins are taken to an inspection station. At may secure the sample with a butterfly net or
the inspection station several methods may scoop as the fruit is being unloaded directly
be used to collect a random sub-sample of the into the plant in the water flume. Still other
Apples - Ch.24 11/4/03 11:02 am Page 629
Fig. 24.3. At a substation, fruit are selected randomly for measuring firmness, Brix and starch to determine
potential storage life.
(a)
(b)
Fig. 24.4. Each bin on the truck is tagged with the grower information (a). The individual tag shows a
bar-code, cultivar, starch reading, Brix and fruit firmness (b).
Apples - Ch.24 11/4/03 11:02 am Page 630
Fig. 24.5. An inspector collects a sub-sample of fruit to be used to determine fruit grade for payment.
companies collect a sample by hand by dig- for the presence of insect larvae during the
ging into the bin to remove fruit (Fig. 24.5). initial inspection.
The subs-ample that is collected is then The remaining apples (US No. 1 grade) are
examined by an inspector (Fig. 24.6). The graded into sizes based upon the minimum
individual is usually hired by an indepen- diameter of the fruit, regardless of its stem or
dent organization run by federal and state calyx position (Fig. 24.7). The following sizes
government, growers and processors. The and criteria are used as outlined in the USDA
inspector is paid by the processing company. Inspection Guidelines: (i) under 2 in. (under
Apples are first inspected and all culls and 57 mm); (ii) 2–2 in. (57–64 mm); (iii) 2–3 in.
juice apples are removed from the sample (64–75 mm); and (iv) 3 in. and up (≥ 76 mm).
and weighed, all weights being recorded Fruit in each size category is then weighed.
to the nearest whole pound (i.e. 0.45 kg). The percentages of the different grades and
Depending upon the processor and the sizes are calculated on the basis of the total
supply and demand for apples, apples in the weight of the sample. These percentages are
US No. 2 grade may also be removed and then recorded and applied to the weight of
weighed at this time. Apples are examined the entire truck-load to determine the return
Fig. 24.6. An initial inspection of fruit done where ciders, culls and No. 2 grades are removed.
Apples - Ch.24 11/4/03 11:02 am Page 631
of these holes has been enough for process-

ing companies to reject entire loads of fruit.
However, damage by surface-feeding insects
that do not penetrate into the fruit is graded
on trim waste. Studies have shown that
apples damaged by surface-feeding insects,
such as tufted apple bud-moth (Platynota
idaeusalis (Walker)), lost more weight in stor-
age and did not store as well as uninjured
fruit (Hull and Rajotte, 1988). Since the
grower is paid on the initial weight at the
time of inspection, the loss in weight and
shorter storage life are an unrecoverable loss
to the processing company. Bruising is scored
on a trim-waste basis and varies depending
Fig. 24.7. At the final inspection the remaining on grade requirements. As mentioned previ-
No. 1 grades are graded into sizes and the ously, growers must judiciously work with
percentage in each is determined and is used to their harvest crews to minimize the amount
determine payment to the grower. of bruising that occurs during the harvest.
Firmness is an important attribute that
the grower will receive. The payment for the determines the potential storage life, as well
apples is based on the size, grade and quality, as the possible end-product. The higher-
at the discretion of the buyer. value products require the firmest fruit.
The inspector is also responsible for cor-
rectly identifying the cultivar that is deliv-
24.10 Grade Interpretations for ered. In the case of mixed cultivars, a lot
Payment may be certified as such when designated by
the processor/grower contract. When lots
The USDA Standards for Grades of Apples are not designated as ‘mixed cultivar’ and do
for Processing in the USA are based primar- contain two or more cultivars, the inspection
ily on the amount of ‘trim waste’ occurring is based on predominant cultivar.
during ‘the usual commercial preparation for The USDA grade standards do not set the
use’. ‘Commercial preparation’ presumes the price for the fruit, and individual processing
use of power-peeling machines, followed by companies may have more or fewer price
any necessary hand-trimming. Trim waste is categories. Some processors have developed
described as the amount of the fruit that their own requirements, based upon their
would need to be removed due to damage or needs. Processors that make products other
decay. The inspector must use considerable than juice realize the importance of fruit size
judgement as to the percentage of fruit flesh and its impact upon the efficiency of the
that needs to be removed (Anon., 1986). As a peeling operation. Individual processing
general policy, in removing bruises or shal- companies may impose additional standards
low defects, a rounded or curved cut should on the fruit. The processors may pay an
be used. For injuries that penetrate deep into additional premium for fruit that meet
the flesh, a cone-shaped cut should be minimum standards, as a means of enticing
employed. US No. 1 and US No. 2 grades growers to produce fruit of the quality
differ from each other based upon the per- desired by the processor.
centage of trim waste. Primary components
that affect trim waste include the amount of
bruising, fruit decay and insect damage. 24.11 Future Directions of the Industry
Apples containing any insect larvae or
holes indicating larvae entry are not allowed A greater emphasis on delivering large bruise-
in any grades. In recent years, the presence free fruit will become more important in the
Apples - Ch.24 11/4/03 11:02 am Page 632
future. The ability to store fruit for 12 months The reduction in trade barriers will
will mean that fruit coming out of storage increase the world movement of apple prod-
must be of sufficient quality to be utilized for ucts, particularly AJC. The development of
any product that is needed. Processing compa- free-market economies in eastern Europe
nies cannot afford to store fruit for an and western Asia have the potential to
extended period of time if they come out of further increase the production of AJC.
storage suitable only for making juice. Unfortunately, in the short run, the over-
The development of preharvest and production of AJC from these areas will only
storage technologies such as the use of serve further to depress world apple prices.
aminoethoxyvinylglycine (AVG) and 1- Initial entry into the world markets of AJC
methylcyclopropene (MCP) will provide the from countries such as the PRC have had an
ability to increase the number of cultivars for impact on world AJC prices. In the future,
processing use. Cultivars that previously had higher-value products, such as apple sauce
a short storage life may now be utilized or sliced apples, may enter world trade. The
due to increased storability. Current cultivars world demand, however, may limit the
utilized can come out of storage with higher quantity that can be profitably sold. Rising
quality and provide greater flexibility in how energy costs will have a direct impact upon
the fruit may be utilized. Processing com- the trade and profitability of processed prod-
panies will need to provide sufficient incen- ucts. A budget analysis indicated that trans-
tives for growers to produce cultivars and portation costs is a major factor that affects
grades of fruit that are most profitable for interregional competition within a county
both the orchardist and the processor. (Jordan, 1983). We can assume that world
Undoubtedly, there will be an even greater markets will be similarly affected by trans-
emphasis on reducing pesticide applications portation costs between different countries.
for the production of processing apples.
There is potential for the use of apple-scab
(Venturia inaequalis (Cooke) Wint.)-resistant Acknowledgements
cultivars. Greater emphasis on the quality of
fresh fruit will mean that growers will no Appreciation is expressed to Mr David
longer be economically able to divert low- Cox, Knouse Foods Cooperative Inc., and
quality fresh-market apples into the process- Mr David Benner, El Vista Orchards, for
ing market as a cost-effective technique. The background information and Knouse Foods
quality demands for high-quality processing Cooperative Inc. for permission to photo-
and for fresh-market apples are becoming so graph the fruit-receiving and grading
different that fruit will no longer be able to be process. The photographs were taken by Dr
diverted from one outlet to the other. George M. Greene, II.
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Apples - Ch.24 11/4/03 11:02 am Page 634
Apples - Index 11/4/03 11:02 am Page 635
Index
abscisic acid 439, 440 aminoethoxyvinylglycine (AVG) 442

abscission, role of ethylene 440 inhibition of ethylene biosynthesis 441, 455
ACC see 1-amino-cyclopropane-1-carboxylic acid role in branch formation 441
Accel® 417, 420, 440, 441, 442 role in delaying fruit ripening 441, 455, 589
adjuvants, use with thinning agents 421–422 role in fruit set 417, 441, 450–451
Adoxophyes orana granulovirus 495 role in preharvest drop 163, 441, 451, 452, 632
advection 524 role in vegetative growth 441
advective frost 241, 522, 531, 534, 538 use in bloom delay 537
‘African Carmine’ 42 ammonium thiosulphate 227, 415
AGRIOS 543, 544, 545 ‘Amorosa’ 42
‘Ahra’ 38, 562 amplification restriction fragment length polymorphism,
‘Ahrina’ 38 use in cultivar identification 44
‘Ahrista’ 38, 79, 562 ‘Anabela’ 36
air flow, in relation to freeze avoidance 522, 523 ancymidol, as an inhibitor of gibberellin biosynthesis 439
‘Akagi’ 40 ‘Angold’ 37, 561, 564
‘Akane’ 40, 72 anther culture 33
‘Akibea’ 40 anthers 158
‘Akita Gold’ 40 anthocyanin synthesis 208, 211, 212, 232
Alar anthracnose canker (Pezicula malicorticis)
impact on flowering 156 symptoms and control measures 461
impact on maturation 163 see also 66, 75, 607
‘Alice’ 42 ‘Antonovka’ 45, 99, 126, 221, 222, 223, 621
‘Alka’ 41 ‘Aori 9’ 39
allergenicity apetalous form 44
selection in breeding programmes 47 Aphelinus mali 513, 546, 574
alleyways aphids 491, 512–513
groundcover management 307, 309, 310 apical dominance 439, 440
‘Alps Otome’ 40 Apogee® 442
Alternaria blotch and rot (Alternaria mali = Alternaria alter- application rates and timing 446–447, 448
nata) impact on fruit set 421, 447, 450
symptoms on fruit and leaves 461, 475–476 use for controlling fire blight 447–448
see also 63, 66, 68, 70, 75 use to retard vegetative growth 443
alternate bearing see biennial bearing apomictic seedlings, use as rootstocks 93, 127
aluminium 247, 251 apomixis 2–6, 11, 33
‘Alwa’ 41, 222 apple aphid (Aphis pomi)
Amblyseius andersoni 510 damage symptoms and life cycle 513
Amblyseius fallacis 510 see also 490, 578
‘Ambrosia’ 82 apple bagging 67
American brown rot see brown-rot diseases apple black spot see apple scab
American fruit fly see fruit flies apple blister bark 482, 483
American hawthorne rust (Gymnosporangium globosum) apple-blossom weevil (Anthonomus pomorum)
464 control measures 503, 575, 578
1-amino-cyclopropane-1-carboxylic acid (ACC) 52, 440, damage symptoms 502, 571
597 ecology 502–50, 558
635
636 Index
apple chat fruit 482 apple scab resistance, as a breeding objective, 34, 35, 36,
apple chlorotic leafspot disease 479, 480 37, 38, 39, 41, 43, 44, 470, 560
apple chlorotic leafspot trichovirus (ACLSV) apple scar skin 483
diagnostic methods 480 apple scar skin viroid (ASSVd)
genetic control of resistance 39 cause of apple dapple apple 483
see also 479–480, 484, 485 see also 63, 66, 68, 69, 70, 78, 79
apple dapple apple 483 apple shape
apple dead spur modification with benzyladenine 420, 441
diagnostic method, symptoms and means of trans- modification with Promalin® 162, 453
mission 485 apple stem grooving see apple decline
see also 64, 484 apple stem grooving capillovirus (ASGV)
apple decline 479 diagnostic methods and symptoms 480
apple decline (on ‘Virginia Crab’) 479, 480 see also 484, 485
apple dimple fruit 483 apple stem pitting 479, 481
apple dimple fruit viroid (ADFVd) 483 apple stem pitting foveavirus (ASPV)
apple false sting 484 diagnostic methods 481
apple flat apple genetic control of resistance 39
diagnostic methods, symptoms, host species and see also 484, 485
transmission vector 479, 480 apple union necrosis 479, 481
see also cherry rasp-leaf nepovirus (CRLV) ‘Applethorpe Earlidel’ 36
apple flat limb 484, 485 ‘Applethorpe Summerdel’ 36
apple freckle scurf 484 AR rootstock series 103
apple fruit crinkle 68, 483 archetypes, of cultivars 563, 564
apple fruit crinkle viroid (AFCVd) 482, 483 ‘Ariwa’ 43, 66, 561, 564
apple green crinkle 68, 484 ‘Arkcharm’ 34
apple horseshoe wound 484 ‘Arlet’ 43, 66, 561, 564
apple internal bark necrosis 64, 484 Armillaria root rot (Armillaria mellea) 461
apple juice concentrate (AJC) Armillaria spp., susceptibility of rootstocks 118
pasteurization 618 ‘Aroma’ 42
production in different countries 616, 618, 632 ‘Aromat de vara’ 42
volumes in world trade 616, 621 ascorbic acid see vitamin C concentration
apple leaf pucker 484 ATS see ammonium thiosulphate
apple maggot fly (Rhagoletis pomonella) attract and kill 498
control measures and phenology models 507 auxins 438–439
see also 64, 66, 68, 490 impact on growth 96, 321, 326
apple mosaic treatment of cuttings 132
symptoms 481 AVG see aminoethoxyvinylglycine
see also 63, 66, 70, 479
apple mosaic (Tulare) 479
apple mosaic ilarvirus (ApMV) 481 BA see benzyladenine
apple proliferation 484 Bacillus thuringiensis (Bt) (pesticide) 495, 576, 579
apple pustule canker 484 backcrossing 32
apple ring rot (Botryosphaeria berengeriana) 461 background colour, as a maturity index 589, 592
apple ring russet 484 bacterial diseases 460–467
apple ring-line pattern 484 bark beetle 491, 580
apple ringspot 484 bark measles see internal bark necrosis
apple rosette 484 ‘Baronesa’ 36
apple rough skin 484 Basamid 254
apple rubbery wood 484–485 basitonic growth habit 328
apple rust mite (Asculus schlechtendali) 64, 511, 575 ‘Baujade’ 38, 66, 69, 562
apple sauce 619–620 bearing habits of specific cultivars 62, 65, 67, 68, 70, 71,
apple sawfly (Hoplocampa testudinea) 503, 571, 575, 578, 72–73, 74, 76, 77, 78, 80, 156, 157, 328
579 Beauvaria bassiana (microbial insecticide) 495
apple scab (Venturia inaequalis), disease characteristics bed systems 204, 206, 386
control measures 461, 470, 573 bees 159
epidemiology 468–469 ‘Bellida’ 41
infection periods 470 ‘Belmac’ 35, 75
influence of leaf wetness 470 ‘Ben Hur’ 77
influence of temperature 469, 470 bench grafting 140, 143
potential ascospore density 470 benzimidazole fungicides 594
resistant cultivars 35, 36, 66, 68, 70, 75, 79, 560, benzyladenine (BA)
561 chemical formula 439
susceptibility of rootstocks 118 impact on feathering of young trees 144, 145, 441, 445
symptoms 461, 468 role in flower bud formation 441
see also 557, 571 role in fruit size, shape and firmness 420, 441
apple scab, genetics use as a thinning agent 411, 417, 420–421, 441
bacterial artificial chromosome library 49 see also 440, 442
genetic control 44 6-benzyladenine see benzyladenine
molecular maps 49 6-benzylaminopurine, role in budbreak 225
molecular markers 47 Bertha army worm (Mamestra configurata) 509
sources of resistance 44, 45 biennial bearing
Index 637
impact of root pruning 336 chilling requirement 77

in relation to organic production 560, 562 cold tolerance 221, 222
in young trees 392 controlled atmosphere requirements 78, 596, 607
see also 72, 155, 162, 210, 328, 410, 411, 413, 428–429, disease and pest susceptibility 78, 470, 561
448 fruit characteristics 77
bins, sanitation 608 global distribution 77
biodynamic production systems 553 global production 18
biological control 500–501, 608 origin and parentage 77
biological diversity, in IFP systems 541 scald susceptibility 77
bitter pit sports 78
control measures 594, 605 storage requirements 77–78, 596
cultivar susceptibility 66, 67, 72, 605 uses, as fresh and for processed products 77
impact of harvest date 587, 605 see also 9, 564, 578, 592, 605
impact of magnesium 290 ‘Bramley’s Seedling’, storage requirements 596
impact of nitrogen 283 branch bending
impact of water stress 178 impact on flowering, hormone concentrations and
treatment with calcium 594, 605 shoot formation 337–338
bitter rot (Glomerella cingulata, Colletotrichum acutatum) methods of 338, 339
461, 473 branching
see also 68, 462, 607 stimulation by notching 341, 445
black pox (Helminthosporium populosum) 476 stimulation with Promalin® 444, 445
see also 461 breeding
black root rot (Xylaria mali) 461 objectives
black rot (Botryosphaeria obtusa) 461, 473–474, 607 apple scab resistance 34, 35, 36, 37, 38, 39, 42, 43, 44,
black spot see apple scab 470, 560
‘blackheart’ injury 220 fire blight resistance 45–46
‘Blair’ 35 fruit quality 34, 36, 38, 39, 40, 42, 46, 47
blister bark (Pseudomonas syringae) 460, 466, 571 growth habit 34, 39, 42, 43
blister canker (Biscogniauxia marginata) 461 pest tolerance 35, 38
blister spot (Pseudomonas syringae) 460, 466 powdery mildew resistance 34, 36, 39, 42, 43
block cutworm (Agrotis ipsilon) 509 storage life 35, 40, 42
bloom winter hardiness 35, 42
date, relationship to timing of harvest 589, 592 programmes in different countries 34–43
delay bridge grafting 241
chemicals for 537 ‘Brina’ 39
for freeze avoidance 536–538 Brooks fruit spot (Mycosphaerella pomi) 462, 475
use of evaporative cooling 226, 537 brown core 605, 606
use of reflectants 537–538 brown-rot blossom blight 462
use of shading 538 see also blossom blight
blossom blight (Monilinia laxa) 462, 472, 572 brown-rot diseases 461, 472, 571 see also Monilinia
blotch (Phyllosticta solitaria) 462 bruising
blue mould (Penicillium spp.) causes 586, 593
control measures 462, 478, 607–608 impact on processing grade 625, 627, 631
symptoms 462, 478 bud temperature, relationship to air temperature 225
‘Blushing Golden’ 72 bud wood, sources, preparation and storage 141
‘Bonza’ 72 budded rootstocks 262
boron budding height 260
availability in soils 246, 293 for organic orchards 558
concentration in irrigation water 293 impact on vigour 98, 141
content in organic matter 293 budding methods 140, 141, 142
critical leaf concentrations 270 buds, low temperature thresholds 223, 532
critical fruit concentrations 270 Bull’s eye rot (Pezicula malicorticis) 66, 75, 461, 607
deficiency symptoms 293–294 burr-knots 261
deficiency thresholds 270, 294 fire blight susceptibility 260
role in metabolism 293 impact of rootstock clone 107, 131
supply bush tree system 347
as fertilizers, by fertigation and in organic amend-
ments 294
as foliar sprays 277, 294 CA storage see controlled atmosphere storage
toxicity symptoms 294 ‘Cacanska Pozna’ 43, 72
toxicity thresholds 270, 294 ‘Cadel’ 43, 72
see also 267 CAJ see concentrated apple juice
’Boskoop’ 561, 562, 564 calcium
bot rot (Botryosphaeria dothidea) 68, 465, 473–474, 607 accumulation in fruit 178, 291–292, 322, 331, 411
botanicals, use as pesticides 492 availability in soils 246, 290–291
boundary layer resistance in orchards 174–175, 183 chloride
bourse shoots, responses to light 202 postharvest application 292, 594
‘Braeburn’ use as foliar spray 292
bearing habit and tree form 77, 328 content in fruit 270–271
breeding, use as a parent 41, 78 content in leaves 268–270
browning disorder (BBD) 77–78 content in tree framework and in roots 268
638 Index
calcium continued training method 325, 339, 348, 349–350

critical fruit concentrations 270, 292 use of semi-vigorous rootstocks 348
critical leaf concentrations 270 yield comparisons 381–382
deficiency symptoms 291 yields 256
impact of pruning 322, 331 ‘Chantecler’ 38
mobility in tissues 291 ‘Chanteline’ 38
nitrate, use as a foliar fertilizer 292 ‘Charlotte’ 37
role in metabolism 291 chemical thinning, effect of temperature 423, 426, 428
role of spur leaves 201 see also fruit thinning
supply cherry rasp-leaf nepovirus (CRLV) 480
as foliar sprays 277, 292 chill units 155, 224–225, 239
for pH adjustment (liming) 250 chilling injury, in storage 595, 597
treatment for bitter pit 288–289, 594 chilling requirements
see also 267 cultivar differences 224, 239
calcium-related disorders 291, 594 influence of warm temperature periods 155, 224–225
calendar date as a maturity index 589, 592 models for prediction of 155, 239
calyx 158 of low chill cultivars 155, 217, 224, 239
calyx-end rot (Sclerotinia sclerotiorum) 462 of specific cultivars 63, 65, 67, 68, 70, 73, 74, 76, 77, 78,
‘Cameo’ 33 80, 155
canopy form substitution with chemicals 217, 225, 239
impact on light interception 184, 197, 199, 200, 204, chimeras 43
377, 385–391 ‘Chinatsu’ 40
in relation to freeze protection 534 ‘Chinook’ 35, 70
canopy shape chip budding 141, 142
conical 348–365 chitin synthesis inhibitors 505
flat fan 365–371 chlorinated hydrocarbons, use as pesticides 492
spherical 347–348 chlorinated sulphur acaricides 496
V 371–378 chlorine 267, 268
Y 204, 206, 207, 372, 381–382, 395, 396–399 chlormequat
captan impact on flowering 156
use in fruit thinning 415, 416 use in bloom delay 537
use as postharvest treatment 608 2-chloroethylphosphonic acid see ethephon
carbamate pesticides 490, 494 chlorophyll degradation 208, 211, 600, 601
carbaryl chlorophyll fluorescence 178
impact on fruit injury 418–419 chloropicrin 254
impact of shade on effectiveness 426 chromosome numbers 2–6, 9, 157
impact of temperature on effectiveness 227 ‘Chukwang’ 40
use in IPM systems 419 cider apples
use as a thinning agent 411, 417, 418–419, 421, 428 cultivar types 618–619
see also 494 production areas 618
carbohydrates rootstock selection 111, 618
role in cold acclimation 223–224 understorey management 308, 618, 627
role in flowering 209 classification of species
carbon dioxide concentration based on intergeneric crossability 9
effect on ethylene production in fruit 601 based on molecular polymorphisms 9, 12
effect on fruit respiration 600 using chromosome number 9
carbon dioxide injury 607 using flavonoids 11, 12
carbon partitioning 201–202 using morphological traits 9, 11
‘Caricia’ 36 clay powders, for disease control 573, 577
carotenoids 208 clearwing moths 580
carpels 157, 161 climacteric 163, 440, 586
‘Catarina’ 36, 68 Clitocybe root rot (Armillaria tabescens) 462
CCC see chlormequat clonal rootstocks see rootstocks
cedar apple rust (Gymnosporangium juniperi-virginianae) cloning genes 52
464 clothes-pegs, for manipulating crotch angles 326, 341,
cell 349, 352, 355, 361, 367, 374, 376
division CO2 assimilation rates see photosynthesis
duration in fruit 161, 201, 229 coconut soap, use as a fungicide 573, 578
impact on fruit size 413 Codex Alimentarius 554
impact of photosynthesis 201 codling moth (Cydia pomonella)
impact of temperature 161, 229–230 biological control 495, 505, 576, 580
impact of water deficit 176, 177 chemical control 504–505
role of cytokinins 321, 440 control using mating disruption 505–506
expansion control using sterile insects 491, 506
impact of water deficit 176 damage symptoms 504
in fruit 161, 229 detection methods 504
central leader system development model 500
light distribution within canopies 388–390 geographic distribution 503–504
planting densities 348 granulovirus 495, 506, 579
profitability 396–399 host range and life cycle 504
support system 380 use of pheromone traps 504
Index 639
see also 66, 489, 490, 491 ‘Cox’s Orange Pippin’

cold hardiness bearing habit 80, 328
impact of cultural practices 223–224, 324–325 breeding, use as a parent 33, 69, 81
influence of mineral nutrients 224 chilling requirement 80
of specific cultivars 62–63, 65, 68, 70, 71, 73, 74, 75, 76, clone T12 81
78, 80, 221, 222 cold hardiness 80, 222
tissue sensitivity 218 controlled atmosphere requirements 80, 596
use of interstems 139 Cox’s disease 81
variation amongst rootstocks 99, 218, 221, 222 disease susceptibility 81, 141, 561, 562
cold tolerance, rootstock breeding for 99 frost susceptibility 80, 227
collar rot resistance fruit characteristics 80
of interstems 139 global distribution 80
of rootstocks 95, 99, 104–105, 108–110, 111, 112, global production 18
114–115, 116, 118, 141 insect pest susceptibility 66, 81
colour charts, for measuring background colour 592 length of growing season 228
coloured traps, for insect control 579–580 origin and parentage 80
columnar cultivars 35, 37, 75 selection of rootstock 96
columnar growth habit, molecular markers 48 sports 81
compensation point 198 storage requirements and disorders 80, 596
concentrated apple juice (CAJ) see apple juice concentrate time of bloom 228
(AJC) tree form 80
condensation (of water) 529 uses, as fresh and for processed products 80
‘Condessa’ 36 yield, relationship to temperature 231
conduction see also 564, 578, 588
definition 524 crab apples
see also 240, 525, 527 as ornamental species 35, 38
conic shaped canopy types as sources of pollen 159, 260
central-leader system 348, 349 ‘Creston’ 35, 66
HYTEC system 358–362 ‘Cripps Pink’ see ‘Pink Lady®’
meadow orchard systems 363 ‘Cripps Red’ 36
mini-central-leader system 348, 350 critical leaf and fruit nutrient concentrations 270
North Holland spindle system 353 critical temperatures, for flower buds 226, 531, 532
palmette-leader system 350–351 crop coefficient, for estimating evapotranspiration
performance characteristics 363–365 183–184
slender-pyramid system 357–358, 359 crop load
slender-spindle system 351, 352, 353 impact on fruit size and maturity 162
SolAxe system 356–357
optimal values 411
super-spindle system 362–363
crop water deficit index 171
vertical-axis system 354, 355–356
cross pollination 157
consumer demand 23, 25, 26
crotch angle
consumption, per capita 23–25
impact of notching 341
controlled atmosphere (CA) storage
impact of pruning 321–322, 328
application of rapid CA 602
see also 460
effects on ethylene production 600
crown gall (Agrobacterium tumefaciens) 466
effects on metabolism and respiration 600
infection of rootstocks 118
impacts on marketing strategies 19
recommendations for low oxygen CA 603 see also 101, 126, 460
recommendations for relative humidity 600 crown rot see Phytophthora
recommendations for standard CA 602–603 crown rot resistance 63
controlled atmosphere requirements, of specific cultivars of rootstocks 95
63, 65, 67, 69, 70, 71–72, 73, 75, 76, 78, 79, 80, 596, cryolite, use as a pesticide 492
607 Cultar® see paclobutrazol
convection 527 cultivar
definition 524 identification, use of microsatellite markers 48
cool chain management 597, 598 selection
copper for cider production 618–619
availability in soils 246, 298 for fresh market supply and for processing
critical leaf concentrations 270 255
deficiency symptoms 298 for organic production 255, 560–562
role in metabolism 298 types for cider production 618–619
supply as foliar sprays 277, 299 cultivars see individual cultivars by name
see also 267 cultivated apple, centre of origin 1, 92–93, 238
copper chelate, use as a defoliant 146 cultivation of soils 311, 314–315
copper fungicides, for control of apple scab, bark canker cultivators for organic production systems 565–567
and European canker 578 cuttings 94, 127
‘Corail’ 38, 66, 82, 561, 564 cutworms 509
cordon system see super-spindle system cyanamide see hydrogen cyanamide
core flush 232 ‘Cybele Delrouval’ 38
corkspot 67, 72, 283 cyme 157
‘Cortland’ 34, 75, 222, 328, 470, 596, 603 cytokinins 439–440
cover crops, for control of soil-borne diseases and nema- impact on vigour 96
todes 309 role in apical dominance 440
640 Index
cytokinins continued descriptive sensory analysis 35

role in cell division 321, 440 dew point
role in leaf senescence 440 definition 529
measurement 533
see also 530
daily thermal cycle 526–528 Diaporthe canker (Diaporthe tanakae) 462
‘Dalinbel’ (‘DL 11’) 79 diatomaceous earth (use as a pesticide) 497
damage action threshold 305 diffuse irradiation
daminozide characteristics 196
effect on flowering 339 effect on fruit quality 207
use in bloom delay 537 dinitro compounds, use as pesticides 492
DARE see Durable Apple Resistance in Europe dinitro-ortho-cresol (DNOC)
DDD 493 role in bud break 225, 239
DDT 248, 493, 504 role as a thinning agent 413, 415
deficiency nutrient concentrations and symptoms use as a pesticide 492
boron 270, 293–294 diphenylamine (DPA) 65, 67, 69, 75, 76, 593–594, 606, 607,
calcium 270 608
copper 270, 298 Diplodia canker (Botryosphaeria stevensii) 462
iron 270, 296 direct irradiation 526
magnesium 270, 290 ‘Discovery’ 45, 49, 564
manganese 270 disease resistance
nitrogen 270, 283 pyramiding (of resistance genes) 45
phosphorus 270, 287 of rootstocks 118, 257
potassium 270, 288 use of interstems 119
sulphur 270, 293 disease resistant cultivars for organic production 561–562
zinc 270, 295–296 disease susceptibility of specific cultivars 63–64, 65–66
deficit irrigation 185–186 68, 69, 70, 72, 73, 75, 76, 78, 79, 81, 141, 470, 473, 475,
defoliation 476, 561, 562
impact on flowering 224, 239 diuron 248
of nursery trees 146–147 DN-11, use as a pesticide 492
defruiting, impact on vegetative growth 414 DNOC see dinitro-ortho-cresol
degree-day models dock sawfly (Ametastegia glabrata) 509
for bloom prediction 225 dolomite 250
for fruit growth 229 dormancy 224–225
for insect growth stages 500, 504 dormant pruning
for shoot growth 231 effects of delayed pruning 325
for yield prediction 231 timing in relation to freeze injury 324–325
‘Delbard Jubilee’ 38, 66, 561, 564 DPA see diphenylamine
‘Delbarestivale Delcorf ’ (‘Delbard Estivale’) 38, 561, 564 ‘Drakenstein’ 42
‘Delblush’ 38, 66, 82, 561, 564 Drilling system see Mikado and Drilling systems
‘Delcorf ’ 66 drought avoidance 179
‘Delearly’ 38 drought tolerance, influence of rootstock 93, 95, 98, 107,
‘Delgaly’ 69 119, 127, 384
‘Delicious’ ‘Dulmener’ 45
bearing habit 62, 156, 157, 160, 162, 328 ‘Duquesa’ 36
breeding, use as a parent 64, 66, 69 Durable Apple Resistance in Europe (DARE) 45, 49
chilling requirement 63, 68 ‘Durello di Forli’ 45, 49
cold hardiness 62–63, 222 dwarfing selections of rootstocks 103
controlled atmosphere requirements 63, 596
disease susceptibility 63, 470, 475
fruit characteristics 63, 453 earth’s energy balance 525
global distribution 62 earthworms 569
global production 18 earwigs 509
insect pest susceptibility 63–64, 66 East Malling series see Malling series; Malling–Merton
manganese toxicity 247 series
maturity indices 63 Ebro trellis system
origin and parentage 62 light distribution within canopies 390–391
pollination requirements 157–158 planting density and selection of rootstocks 370
pruning requirements 324, 325, 326 pruning and training methods 370
scald susceptibility 63, 593 ecdysone agonists (pesticides) 505
sports 43, 64 ‘Ecolette CPRO’ 41, 79, 562
spur types 64 ecological compensation areas 541, 559–560
storage disorders 63, 605 economic injury level (EIL) 305, 499, 540
storage requirements 63, 596, 597, 603 see also 491, 510
transformed selections 50 economic performance
tree form 62 of different orchard systems 378, 384, 395–400, 401,
uses, as fresh and for processed products 63 402
see also 9, 12, 76, 126, 601–602 sensitivity to fruit price and land prices 398, 399, 400
‘Delios de Voinesti’ 42 sensitivity to yields 398, 399
demand: supply balance 27 effective pollination period (EPP) 159
deposition (of water) 529 ‘Eir’ 41
Index 641
‘Elan’ 41 genetic control 52

elevation, effect on temperature 238, 239, 527–528 impact of controlled atmosphere 600–601
Elgetol see dinitro-ortho-cresol impact of temperature 595, 597
‘Elise’ (‘Roblos’) 41, 72, 81 inhibition with AVG 441, 451, 589
‘Elstar’ promotion with ethephon 588
bearing habit 78, 328, 562 role of IAA 439
breeding, use as a parent 48, 79 inhibition with 1-MCP 594
chilling requirement 78 removal from CA stores 603
cold hardiness 78, 221, 222 role in flower and fruit senescence 440
controlled atmosphere requirements 79, 596 role in fruit abscission 440, 451
Cox’s disease 79 role in fruit ripening 163, 440, 586, 588–589, 590
disease susceptibility 79, 561 role in induction of flowering 440
fruit characteristics 78–79 European brown rot (Monilinia fructigena) 462
global distribution 78 see also 63, 472, 571
global production 18 European canker see Nectria canker
origin 41, 78 European red mite (Panonychus ulmi) 63, 66, 69, 78, 476,
parentage 66, 78, 81 489, 510, 546, 575, 579
scald susceptibility 79 ‘Eva’ 36, 72
selection of rootstock 96 evaporation 529
sports 79–80 evaporative cooling
storage requirements 79, 596 application methods 226
summer pruning 79 timing of application 227
transformed selections 50 use in bloom delay 226, 537
tree form 78 use to prevent sunburn 230
uses, as fresh and for processed products 79 evapotranspiration
see also 33, 64, 383, 564 estimation methods 182–183
embryo 160, 161 impact of environmental factors 182, 184
‘Empire’ impact of ground-cover crops 307
bearing habit 156, 328 influence of stomatal conductance 183
cold hardiness 221, 222 modelling of 182–183, 186
controlled atmosphere requirements 596, 603, 607 ‘Evereste’ 38
disease susceptibility 561 exogenous plant bioregulants 440–443
fruit characteristics 564 see also growth regulators
origin 34 exports
parentage 64, 75 influence of exchange rate 22
scald susceptibility 593 of concentrated apple juice 616, 621
sports 43 of fresh produce 21
storage requirements 596, 603, 607
tree form 329, 330
see also 162, 383 Fadengerät 571
endodormancy see dormancy ‘Fall Red’ 35
endogenous plant hormones 438–440 ‘Falstaff ’ 33, 66
Endothall 415 families and sub-families, of Malus 10
energy content of air and water vapour 528 ‘Fantazja’ 41, 221, 222
‘Enterprise’ 34, 64, 66, 75 feathered trees, pruning management 262, 325, 352, 353,
epidermal cells 161, 210 360
eriophyd mites 511 feathering
espalier tree forms 347, 365 impacts of growth regulators 144–145
ethephon impacts of leaf removal 144
application rates and timing 419, 428, 442, 448, 449, impacts of temperature 143
454, 455 impacts of tipping 143–144
associated use of NAA 454 impacts on yield 143, 258
cultivar responsiveness 419 fermented cider see cider apples
impact on flowering 156 fertigation 275–276, 278, 285, 290
impact on fruit maturation 163, 454, 588 fertilization, of flowers 160
impact of temperature on effectiveness 227, 419, 426 fertilizer management
role in fruit set 420 application of foliar sprays 276–277
role in return bloom 420, 429, 448 application of solid fertilizers 274–275
use to advance ripening 449, 454, 588 application of soluble fertilizers 275–276
use as a defoliant 147 for organic production systems 277–278, 556, 565
use to enhance skin colour 454–455 in intensive systems and with young plantings
use on non-bearing trees 449 395
use to promote flower bud formation 362, 363, 449 field capacity, definition of 245
use as a thinning agent 411, 417, 419–420, 428, 448 ‘Fiesta’ (‘Red Pippin’) 37, 48, 49, 72, 81, 564
see also 440, 441 fire blight (Erwinia amylovora)
Ethrel® see ethephon control measures 447–448, 467
ethylene 439, 440 epidemiology 466–467
as a harvest index 589, 590 host species 466
biosynthesis influence of temperature 466, 467
by fruit 586 inheritance of resistance 46
cultivar differences 455, 586 predictive models 467
642 Index
fire blight (Erwinia amylovora) continued iron 277, 297

resistance manganese 277, 298
as a breeding objective see breeding objectives magnesium 277, 290
of ‘Golden Delicious’ 65 nitrogen 277, 285
of rootstocks 108, 111, 112, 114–115, 116–117, 118, phosphorus 277, 287–288
257, 467 potassium 289
resistant species 46 zinc 277, 296
somaclonal variation 46, 49 forced-air cooling 598, 599
susceptibility, of rootstocks 95, 99, 467 formalin 254
symptoms 460, 466 ‘Fred Hough’ 36
‘Firiki’ 38 ‘free’ palmette system 366, 367, 368, 380
fisheye rot (Butlerelfia eustacei) 462 ‘Freedom’ 34, 66
flat planar canopy types 365–371 freeze
Ebro trellis system 369–370 injury (winter injury)
espalier systems 365 cultivar differences 221
‘free’ palmette system 366–368 dehardening 220, 241
horizontal palmette system 366 impacts of cultural practices 223–224, 241, 284, 325,
Lincoln canopy system 369 531
oblique palmette system 365–366 protection methods 220, 223–224
palmette systems 365–366 recovery from 220, 222, 241
Penn State thin-wall trellis system 368–369 sensitivity of different tissues 222, 225–226, 532
’regular’ palmette system 365–366 significance 218
Solen system 370 species differences 221
tabletop bed system 371 symptoms 220
flavonoids 208 probability
flavour impact of airflow 226
heritability 44 impact of dew-point 533
of fruit 43, 586–588 influence of ground cover 303
see also breeding objectives role of soil type 522, 523
flavour groups, of cultivars 563, 564 protection methods
flesh browning, selection in breeding programmes 47 bloom delay 226–227, 536–538
flesh colour cultivar selection 226
importance in processing 624 heaters 226, 527, 533–534, 536, 538
use as a harvest index 589, 592 overhead irrigation 226, 534–535, 538
flesh firmness, as a maturity index 589, 590 reflectants 531
‘Florina’ 33, 38, 64, 66, 561, 564 site selection 218, 239
flower bud development under tree irrigation 535, 538
models for prediction of 225 wind machines 226, 240, 527, 528, 535–536, 538
frost susceptibility 226, 531, 532 temperatures
role of temperature 225 forecasting 531, 533
flower bud differentiation, influence of light 208, 209, 210 measurement of 531–532
flower bud inhibition, role of gibberellins 155, 209, tolerance
449–450 of cultivars and rootstocks 221, 222, 241
flower induction 154 of interstems 139
flower initiation 155, 156 types 240–241, 531
flower thinning 414, 571, 572 fresh market fruit
flowering cultivar selection 18, 19, 255
genetic control 52 distribution systems 25–26
habit, of axillary buds and terminal buds 157 trade patterns 20–22
impact of growth regulators 155, 156, 209, 449–450 standards 410
impact of heavy cropping 155 frogeye leaf spot (Botryosphaeria obtusa) 461, 474
impact of nitrogen 156 frost point 529, 530
impact of pruning 428–429 frost susceptibility of specific cultivars 62–63, 65, 67, 71,
impact of ringing and scoring 156 72, 73, 74, 76, 80, 227
impact of rootstock 96 frost tolerance thresholds, for developing flower buds
impact of shade 156 226, 523
impact of summer pruning 333 fruit abscission
impacts of branch bending 156, 337–338 influence of growth regulators 163
in tropical climates 155 influence of shade 209, 210
juvenile period 32, 153 role of ethylene 440, 451
flowers 157, 158, 225–226 role of IAA 439
flyspeck (Schizothyrium pomi) 462 fruit acidity, molecular markers 48
causal organism 475 fruit bagging, impacts on colour and fruit finish 210–211
control measures 475, 578 fruit characteristics of specific cultivars
symptoms 474 ’Braeburn’ 77
foliage feeding insects 509–512 ‘Cox’s Orange Pippin’ 80
foliar fertilizers, use in organic production 570 ‘Delicious’ 63, 453
foliar nutrient sprays ‘Elstar’ 78–79
boron 227, 294 ‘Empire’ 564
calcium 292 ‘Fuji’ 67, 208, 210
copper 277, 299 ‘Gala’ 70, 410
Index 643
‘Golden Delicious’ 65, 622 influence of light intensity 160–161, 208, 209, 424–426,
‘Granny Smith’ 69, 162 427
‘Jonagold’ 73, 410 influence of nitrogen 161
‘Jonathan’ 71, 410 influence of pruning 161, 428–429
‘McIntosh’ 74, 232, 413 influence of rootstock 96, 161
‘Rome Beauty’ 76, 622 influence of temperature 161, 428
fruit colour promotion with ethephon 420
effect of light intensity 211, 391 role of IAA 439
effect of summer pruning 72, 331, 332, 333, 334 fruit shape
molecular markers 48 impacts of temperature 162
fruit damage modification using Promalin® 162, 453
caused by insects 501–509 role of chemical thinners 413
caused by pollenicides 415 fruit size
fruit disorders, impact of water stress 177–178 effect of benzyladenine 420
fruit distribution (within canopies), influence of light 210, effect of scoring 340
387–391 effect of temperature 229
fruit drop 160, 201, 441, 451, 455 effect of thinning time 162
fruit firmness effect of V-shaped canopies 378, 381
importance in processing grades 624 heritability 44
use as a harvest index 589, 590, 592 impact of crop load 162
fruit flavour and aroma 44 impact of flower position 162
fruit flies (Anastrepha fraterculus, Ceratitis capitata) 506–507 impact of root pruning 336
fruit growth impact of rootstock 162, 257
impact of water stress 185 impact of seed number 162
response to shade 424–426 impact of shade 162
response to temperature 229–230 impact of spur leaf area 162
role of IAA 439 impact of thinning 411, 412, 414
seasonal growth pattern 185, 229 in relation to time of harvest 161, 164, 589
fruit maturation prediction 162
cultivar differences 162 fruit solute potential 169, 179
impact of cropping density 162 fruit structure 158
impact of growth regulators 163, 441, 455 fruit texture
impact of rootstock 162 quantitative trait loci 46
impact of solar radiation 163 see also 588
impact of temperature 162–163, 229–230 fruit thinning
fruit nutrient concentrations for organic production systems 571, 572
sampling method 270 impact on biennial bearing 161, 410
values for specific elements 268, 270–271, 569 impact on flower bud differentiation 161, 411, 449
within-fruit gradients 271, 291 impact on fruit shape 413
fruit processing impact on fruit size 161, 163, 411
grade standards 410, 620, 624–625, 627–631 impact on vegetative growth 414
payment methods 631 impact of weather 415
quality assessment 627 in relation to insect damage 413
storage methods 625 mechanical methods for 429, 571
fruit quality role of NAA 227, 417–418
as a breeding objective 46 timing 161, 413
effect of canopy form 210, 381, 387, 388, 390, 391 fruiting habit 328
effect of fruit size 411 fruiting zones
effect of light 210 classification 356
effect of summer pruning 322, 331, 332, 333 see also 327
effect of thinning 161, 163, 411 ‘Frumos de Voinesti’ 42
effect of water stress 305 ‘Fu Shuai’ 66
impact of ground cover 311 ‘Fuji’
impact of harvest date 587–588 bearing habit 67, 155, 160, 328, 410, 411
impact of root pruning 336 breeding, use as a parent 68
impact of scoring 340 chilling requirement 67, 68
impacts of temperature 230, 232 cold hardiness 63, 221, 222
selection for 34, 36, 38, 39, 40, 42, 46, 47 controlled atmosphere requirements 67, 596, 607
use of sensory testing 46 disease susceptibility 68, 561
fruit ripening frost susceptibility 67
inhibition with 1-MCP 594 fruit bagging 210–211
role of ethylene 586 fruit characteristics 67, 208, 210
role of IAA 439 global distribution 66–67
use of ethephon 454–455 global production 18
fruit rots (Monilinia fructicola, Monilinia fructigena) 461, insect pest susceptibility 66, 68
472, 473 origin 40, 66
fruit russet see russet parentage 64, 66
fruit set 160 scald susceptibility 67
effect of Apogee® 447, 450 sports 43, 68
effect of AVG 161, 450–451 storage disorders 67, 604
effect of paclobutrazol 451 storage requirements 67, 596
644 Index
‘Fuji’ continued impact on vigour 96

transformed selections 52 use to control of biennial bearing 416, 439
tree form 67 use to control flowering 239, 414, 449–450
use, as fresh and for processed products 67, 620, 621 use to control russeting 453
see also 9, 15, 17, 33, 77, 564, 592 use to stimulate stem elongation 439
functional equilibrium 320 ‘Ginger Gold’ 33, 45, 470
fungal diseases 468–478 ‘Giongo’ 39
fungi, asexual forms (imperfect stage) and sexual forms ‘Giotto’ 39
(perfect stage) 468 Glomerella leaf spot 462
fungicides ‘Gloster’ 18, 564
postharvest applications 607–608 ‘Gold Gift’ 42
see also fungal diseases ‘Golden Delicious’
fusion (of water) 529 bearing habit 65, 157, 160, 328
breeding, use as a parent 33, 36, 48
chilling requirement 65, 68
GA3 439 cold hardiness 63, 221, 222
role in flower bud inhibition 416 control of fruit russet 453
use as a flower bud inhibitor 156, 449 controlled atmosphere requirements 65, 596
GA4 156, 439, 441 cultivar characteristics 65
GA4+7 disease susceptibility 65–66, 470, 475, 476, 561
role in flower bud inhibition 416 fire blight resistance 65
use to control russet 79, 81, 416 frost susceptibility 65
use as a flower bud inhibitor 156, 449 fruit characteristics 65, 622
use to promote feathering in young trees 144 global distribution 64
see also 81, 442 global production 18
GA7 impact of root pruning 336
use as a flower bud inhibitor 416 insect pest susceptibility 66
see also 439 origin 64, 622
‘Gala’ parentage 64
bearing habit 70, 157, 328 precocity 154
breeding, use as a parent 41, 48, 70 pruning requirements 325
chilling requirement 70 scald susceptibility 65
cold hardiness 70, 221, 222 sports 43, 66
controlled atmosphere requirements 70, 596 storage disorders 65, 586
disease susceptibility 70, 470, 473, 561 storage requirements 65, 596
fruit characteristics 70, 410 time of bloom 228
fruit quality 331 transformed selections 50
global distribution 69 tree form 65, 622
global production 18 use as an interstem 120
insect pest susceptibility 70 use in processing 65
origin 69 uses, as fresh and for processed products 65, 620, 621,
parentage 64, 66, 69 622
scald susceptibility 593 see also 9, 12, 45, 76, 565, 590, 604
sports 43, 70–71 ‘Golden Delicious 463’ 42
storage requirements 70, 596, 597 ‘Golden Mira’ 39
transformed selections 46, 50 ‘Golden Orange’ 39
tree form 70 ‘Golden Sentinel’ 35, 75
uses, as fresh and for processed products 70, 620 ‘Goldjon’ 72
see also 9, 17, 33, 64, 77, 564, 592 ‘Goldrush’ 34, 45, 66, 82, 561, 564
gas exchange rates see photosynthesis ‘Goro’ 43, 561
gene cloning 52 gracillarid leaf-miners 511
General Agreement on Tariffs and Trade (GATT) 22 grade standards for processing fruit see process grade
‘Generos’ 42 fruit standards
genetic grafting 140–143
control graft-transmissible agents 96
of ethylene production 52 graft-union necrosis virus 312
of flowering 44 ‘Granny Smith’
of plant architecture 44 bearing habit 68, 328
markers, isozymes 47, 49 breeding, use as a parent 69
Geneva Y-trellis system chilling requirement 68
planting density 372 cold hardiness 68, 221, 222
profitability 396–399 controlled atmosphere requirements 69, 596
pruning method 374 disease susceptibility 69, 470
support system 380 fruit characteristics 69, 162
training method 372, 373, 374 global distribution 68
use of dwarfing rootstocks 372 global production 18
‘Gerlinde’ 38, 79, 562 insect pest susceptibility 69
germplasm centres 33 length of growing season 228
gibberellin biosynthesis inhibitors 439 origin and parentage 68
gibberellins 439 scald susceptibility 69, 593
impact on flowering 155, 156, 209, 439 sports 69, 97
Index 645
storage disorders 69 ‘Haralson’ 34, 221, 222

storage requirements 69, 596, 597 hardening 241
transformation 50 hardwood cuttings 127, 131–133
tree form 68 ‘Harmonie Delorina’ 38
uses, as fresh and for processed products 69, 620, 621, harvest indices 163, 589–592
624 harvest period, in relation to storage period 587–589,
see also 9, 15, 564, 578, 592, 604 592–593
granulosis virus, for control of codling moth 495, 506, harvest timing 587–589
576, 579 ‘Hatsuaki’ 40
Graphania mutans 502 ‘Hawkeye’ see ’Delicious’
‘Gravenstein’ 222, 228, 328, 561, 562, 564, 623 Hazard Analysis and Critical Control Point (HACCP) 17
green apple aphid 575, 578 heading cuts
green fruit-worms (Amphipyra pyrimadoides, Lithophane effects of 323, 324, 392, 444
antennata, Orthosia hibisci, O.incerta) 502 use of 322–324, 349, 352, 353, 392
‘Greensleeves’ 46, 50 heat of condensation 526, 533
grey-mould rot (Botrytis cinerea) heat of evaporation 526
control measures 463, 607–608 heat of fusion 525
symptoms 463 heaters
ground colour see background colour for freeze protection 533–534, 538
groundcover use with wind machines 534, 536
management helicopters, use for frost protection 536
impact on organic matter content 304, 309 herbicide
in alleyways 309, 310 residues 248
in integrated production systems 542 use
in organic production systems 555, 565 application methods 313–314
in relation to fruit quality 310 as thinning agents 421
to avoid frost damage 523 for weed control 305, 307, 309
vegetation impact on organic matter content 304, 310
as habitat for beneficial arthropods 312–313 impact on yield 310
selection of grass and legume species 308, 312 in integrated production systems 542
species composition 311–312 selection of type 313, 314
growing heritability of traits 44
degree-days ‘Herma’ 72
for bloom 225 high carbon dioxide concentration, use as a stress treat-
for fruit growth 229 ment 604
for insect growth 500, 504 high carbon dioxide injury 605
symptoms and cultivar susceptibility 607
for shoot growth 231
high volume spraying, for application of thinning agents
for yield prediction 231
422
season
‘Himekami’ 40, 68
duration 228, 239
history
influence of proximity to water mass 238, 239
development of cultivars 7
growth
development in different countries 7, 8–9
habit
development of rootstocks 92
as a breeding objective 34, 39, 42, 43
role of early civilizations 7
classification of types 328–329
‘Hokuto’ 39, 68
cultivar differences 328 ‘Holsteiner Cox’ 81
influence of rootstocks 94, 96, 98 honey bees, in relation to groundcovers 159, 312
spur bearing types 328 ‘Honeycrisp’ 33, 34, 45, 82, 221, 222, 470
tip bearing types 328–329 ‘Hongro’ 40
regulators horizontal palmette system 365–366, 382
for control of branching and suckering 444, 445 hormones 96
for control of ethylene action 594 ‘Huaguan’ 37, 68
for control of feathering 144–145 ‘Huashuai’ 37, 64, 68
for control of fruit set and preharvest drop 450, ‘Hwahong’ 40
451–452 hybrid tree cone orchard system see HYTEC system
for control of vegetative growth 446–448 hybrid trellis 379
effect on flowering 155, 156, 209, 449–450 hydraulic conductivity of rootstocks 181
effect on fruit appearance and shape 452–453, 454 hydraulic resistance 169
effect on ripening 454, 455 hydrocooling, of fruit 598
respiration 228 hydrogen cyanamide, role in budbreak 225, 239
‘Gunma Meigetsu’ 40 HYTEC system
Gütingen V slender spindle-system planting densities 358
planting densities 375 pruning method 359, 360, 361, 362
profitability 396–399 support system 380
training method 375, 376–377 training method 339, 358, 359, 360, 361, 392
use of dwarfing rootstocks 375 use of dwarfing rootstocks 359
yield comparisons 382
HACCP see Hazard Analysis and Critical Control Point

hairy root (Agrobacterium rhizogenes) 466 IAA see indole-3-acetic acid
haploids 33 IBA see indolebutyric acid
646 Index
‘Idared’ internal ethylene concentration (IEC) 589, 590, 592

bearing habit 328 interspecific hybridization 33
global production 18 interstem
self-incompatibility alleles 48 length, impact on vigour 92, 98, 119, 120
use in processing 621, 623 trees
see also 72, 470, 475, 561, 564, 595, 596 production methods 139, 146
‘Iduna’ 43, 564 use for cold hardiness 119
‘Idunn’ 41 use for disease resistance 119
IEC see internal ethylene concentration use for vigour control 97
IFOAM 553 see also 92, 95
IFP see integrated fruit production inversion layer 536
impact injury see bruising definition of 528
‘Imperatiz’ 36 measurement of 532
in vitro propagation see micropropagation IOBC/WPRS 539–541, 543, 545, 546, 547
inarching 241 IPM see integrated pest management
inbreeding depression 32 IPP see integrated plant protection
inclined (angled) leader 360, 361 iron
incompatibility alleles 48 availability in soils 246, 247, 296
indole-3-acetic acid 438–439 critical leaf concentrations 270
indolebutyric acid (IBA) 132, 134, 135, 439, 442 deficiency symptoms 296–297
induced mutations 43 role in metabolism 296
‘Ingrid Marie’ 78, 81 supply as foliar sprays 277, 297
‘Initial’ 38, 70 supply as trunk injections 297
inorganic pesticides 492 see also 267
INRA ‘Baujade’ 38 irradiation, use in breeding 43
INRA Belchard ‘Chantecler’ 38 irrigation
INRA Perpetu ‘Evereste’ 38 management 394, 523
INRA Querina ‘Florina’ 38 scheduling
in-row spacing, tree management 330 estimation of evapotranspiration 182
insect growth regulators (pesticides) 496 use of fruit diameter and trunk diameter 185
insect pest susceptibility of specific cultivars 63–64, 66, use of leaf temperature 185
68, 69, 70, 75, 76, 78, 81 use of midday stem potential 185
insect use of soil moisture monitoring 182
pests ISHS 539, 542, 543
control strategies and methods 499–501 isozymes, use as markers 47, 49
see also individual pests ‘Ivette’ 81
population growth ‘Iwakami’ 40
factors affecting 499
prediction using degree-day models 500, 504
resistance, molecular markers 48 J.9, impact on drought tolerance 95
integrated control 501 ‘James Grieve’ 45, 222, 564
integrated fruit production (IFP) 499, 501, 539–544 Japanese apple rust (Gymnosporangium yamadae) 465
definition 540 ‘Jarka’ 37
in Argentina (case study) 545–546 ‘Jerseymac’ 34, 222, 564
in Europe 543 ‘Jerseyred’ 77
in Italy (case study) 544–545 ‘Jinguang’ 17, 64
in New Zealand (case study) 546–547 Johnny Appleseed (Jonathan Chapman) 8
in USA (case study) 547–548 ‘Jonadel’ 72
integrated pest management (IPM) ‘Jonagold’
definition of 499, 540 bearing habit 72–73, 328
tactics 500 chilling requirement 68, 73
integrated production systems cold hardiness 63, 73, 221, 222
cultivar selection 541 controlled atmosphere requirements 73, 596
definition and principles 540 disease susceptibility 73, 470, 561
ecological compensation areas 541 fruit characteristics 73, 410
fertilizer use 278, 542 global distribution 72
ground cover management, herbicide use and global production 18
mulching 542 origin 34, 73
nutrient cycles and soil fertility standards 540 parentage 66, 72
pesticide selection 542–543 ploidy 73, 157
postharvest considerations 543 sports 43, 73
quality definition 541 storage requirements 73, 596
site selection 541 transformed selections 51
tree training 542 tree form 72–73
internal bark necrosis 247, 298 tree quality, in nursery 137
internal breakdown 210, 283 uses, as fresh and for processed products 73, 621, 624
internal browning 605 see also 9, 33, 64, 79, 383, 564
cultivar susceptibility 67 ‘Jonathan’
impact of climate 606 bearing habit 71, 328
impact of nitrogen 283 breeding, use as a parent 33, 39, 48, 72
symptoms 606 cold hardiness 71, 222
Index 647
controlled atmosphere requirements 71–72 importance in flowering 157

cultivar characteristics 71 latitude, influence on productive limits 196, 217, 238
disease susceptibility 72, 470, 561 layering 94, 129–131
frost susceptibility 71, 72 L/D ratio, of fruit 162, 163
fruit characteristics 71, 410 lead arsenate 248, 491, 492, 504
global distribution 71 leaf abscission, role of IAA 439
global production 18 leaf area duration, relationship to yield 232
impact of root pruning 335, 336 leaf area estimation
impact of summer pruning 332, 333 of canopies 199
origin 71, 623 relation to tree water use 174
parentage 71 leaf area index, impact on light interception 385, 386
pollination requirement 157 leaf feeding insect pests 491, 509–512
scald susceptibility 72 leaf:fruit ratio, relationship to fruit size 413
sports 72 leaf nutrients
storage characteristics and disorders 71–72 seasonal changes in 268
tree form 71 standard sampling protocol 268–269
uses, as fresh and for processed products 71, 621, 623 leaf removal, for control of feathering 144
see also 564 leaf respiration rate, temperature response 228
Jonathan spot 71–72 leaf senescence 439, 440
‘Jonsib’ 45 leaf water potential 169–170
‘Judaine’ 38 leaf-miners 511
‘Judor’ 38 leafrollers 507–509
juice see apple juice concentrate see also 490, 491, 505
juice content lenticel blotch 605
as a breeding objective 46 Leptosphaeria canker 463
heritability 44 Leucostoma canker and dieback 463
‘Julia’ 37, 562, 564 ’Liberty’ 34, 46, 75, 561, 564
June drop 160, 424, 427 light distribution within canopies
‘Jupiter’ 64 conic-shaped canopies 364, 388
‘Jurella’ 38 impact of canopy system 204, 206, 348, 351, 381–382,
juvenile period 32, 153 388–391
juvenility, effect on layer bed performance 130 impact of pruning 207, 330
juvenoids (pesticides) 505 impact of summer pruning 390, 391
impact of training system 204, 206
impact of tree shape 204, 206, 388
‘Kamhong’ 40 impact of tree size 207, 388
‘Kanki’ 40 impact on flowering 209, 387
kaolin clay 230, 497, 577 impact on fruit colour 387, 291
‘Karmijn de Sonnaville’ 81 impact on fruit distribution 209, 390
‘Katja’ 42, 221 impact on fruit quality 207, 391
‘Kidd’s D.8’ see ’Gala’ impact on fruit size 387
‘Kidd’s Orange Red’ 69, 81, 561, 564 impact on yield 207
‘Kim’ 42 importance of limb removal 390, 391
king flowers 157, 415 multiple-row systems 354
king fruit 162 planar canopies 390–391
‘Kinsei’ 40, 66 V-shaped canopies 377–378, 389, 391
‘Kio’ 40 light intensity
‘Kitakami’ 40 effect on fruit abscission 418
‘Kitanosachi’ 39 effect on fruit colour 211
‘Kitaro’ 40 effect on fruit set 201, 208
‘Kizashi’ 40 effect on primary spur leaves 198–199, 208
‘Klara’ 37 light interception
‘Kogetsu’ 40 descriptive models 202–204, 385
‘Kotora’ 40 influence of row direction 200, 202–203
influence of row width 204, 206, 385, 387
influence of tree height 204, 385, 387
labour input measurement methods 203
in different planting systems 365, 371 in multiple row plantings 204, 206, 386
in IP systems 580 in relation to canopy type 184, 197, 199, 200, 204, 377,
in organic production systems 571, 580–581 385–391
Lacanobia subjuncta 509 relationship to yield 195, 197, 202
lacewings 513 light reflecting mulches 211–212
ladybirds 513 see also 208
‘Lady Williams’ 36 light transmission
Ladurner mechanical hoe 565, 566, 567 effect of pruning 207, 330, 390, 391
latent heat effect of root pruning 337
definition 525 ‘Ligol’ 41, 222
see also 226 limb bending see branch bending
lateral branch promotion, using Promalin® 444 limb positioning, to control vigour 392
lateral buds limb-renewal pruning 391
effect on fruit development 162 see also renewal pruning
648 Index
lime sulphur 578 sub clones 93, 258

liming support requirements 100, 257, 351
materials used for pH adjustment 250 tolerance of winter injury 99, 100, 222
in organic production 570 virus status 130
Lincoln canopy system woolly apple aphid sensitivity 100, 385
light distribution within canopies 390–391 yield efficiency 257
planting density 369 M.9 rootstock clones 103, 106, 385
pruning and training methods 339, 369 ‘Macoun’ 34, 75
rootstock selection 369 macrocyclic lactones (pesticides) 495, 505
yield comparisons 381 magnesium
liners 129, 132 availability in soils 246, 289
‘Lodel’ 41 content in fruit 270–271
‘Lobo’ 75, 221, 222 content in leaves 268–270
low chill cultivars 36, 41, 155, 217, 224, 239 content in roots 268
low ethylene CA storage 603, 606 content in tree framework 268
low oxygen CA storage 603 critical leaf concentrations 270, 290
low oxygen concentration deficiency symptoms 290
for control of superficial scald 593, 604, 606 measurement method 289
use as a stress treatment 604 prevention of necrotic leaf spot 81
low oxygen injury 605 role in metabolism 290
symptoms 607 supply
see also 76 as foliar sprays 277, 290
low temperature breakdown 605 fertilizer application rates 290
impact of climate 606 fertilizer forms 250, 290
role of phosphorus 287, 288 impact of pH 290
symptoms 606 see also 267
see also 595, 596, 597 ‘maiden’ trees 138, 140
low volume spraying, for application of thinning agents ‘Maigold’ 43, 66, 561, 562, 564
422 maintenance respiration 228
‘Lustre Elstar’ see ‘Elstar’ ‘Makedoni’ 38
lygus bug (Lygus lineolaris) 312, 503, 579 maleic hydrazide, use in bloom delay 537
lyonetid moths 511 ‘Malling Kent’ 72
lysimeter, for estimation water use 173 Malling series of rootstocks 99, 347
Malling–Merton (MM) series of rootstocks 95, 99, 118
Malus
M&B 25, 105 astrosanguinea 804
impact on feathering 144, 145 source of scab resistance 44
M.25 rootstock 97, 98, 99, 103 baccata jackii
M.26 rootstock source of scab resistance 44
alternative rootstocks with similar vigour 103, 107, classification 2–6
112–113 floribunda 821,
impact on fruit maturation 94 source of scab resistance 45
origin 99 micromalus
propagation 130, 133, 134, 135 source of scab resistance 44
sensitivity to drought 180 related genera 9–11
transformed selections 51, 100, 107–108 species 11–12
use as an interstem 95, 102, 107 angustifolia 2, 12
M.27 rootstock asiatica 7, 12
alternative rootstocks with similar vigour 103, baccata 2, 7, 12, 44, 110, 221
104–105, 256 coronaria 2, 12, 42
impact on drought tolerance 95 dasyphyllus 1
impact on fruit size 94, 257 florentina 2, 12
origin 99 floribunda 3, 7, 12, 45, 48, 77
productivity 204, 205 fusca 3, 12
propagation 128, 133 ioensis 3, 12
root growth 180 mandshurica 3, 7, 12
tolerance of winter injury 99 micromalis 3, 7, 12, 44, 110
M.7 rootstock orientalis 3, 7, 12
suckering 257 praecox 1
transformed selections 51, 100 prattii 3, 12
M.793 rootstock 99 prunifolia 4, 7, 12, 93, 102, 116, 127, 221
M.9 rootstock pumila 4, 12, 92
alternative rootstocks with similar vigour 103, robusta 45, 46, 47
106–107, 108–110 sargentii 4, 7, 12, 93
fire blight susceptibility 100, 385 sieboldii 4, 7, 46, 93
influence on fruit size 94, 257 sieversii 1, 4, 7, 61, 93, 102, 127, 221
influence on scion growth habit 329 sylvestris 1, 4, 7, 12, 42
origin 99 toringoides 4, 12, 93
productivity 204, 205 transitoria 4, 12
propagation 128, 129, 130, 133, 135, 137, 141 tschonoskii 4, 12
sensitivity to drought 180 x asiatica 5
Index 649
x atrosanguinea 5, 7, 44, 47 with V-shaped canopies 378

x domestica 1, 7, 12, 92 pruning, impacts on growth 330, 369
yunnanensis 5, 12 thinning 429
zumi 5, 7, 45, 47 Mediterranean fruit fly (Ceratitis capitata) 507
subfamilies 10 ‘Megumi’ 39, 72
see also 2–6, 12, 46 ‘Melba’ 75, 222
manganese ‘Mellow’ 39
availability in soils 246, 247, 297 ‘Melrose’ 64, 72, 222, 336
critical leaf concentrations 270 ‘Meridian’ 81
deficiency symptoms 298 ‘Merlijn’ 36
prevention of necrotic leaf spot 81 Merton 793 rootstock 111, 116, 118
role in metabolism 298 Merton Immune rootstock series 99
supply as foliar sprays 277, 298 1-methylcyclopropene (1-MCP)
supply as trunk injections 298 use to delay ripening 594, 632
toxicity symptoms 247, 298 MIA trellis system
see also 267 planting density and training method 373
manures, use in organic production 567, 569 use of semi-vigorous rootstocks 373
marigold 254 yield comparisons 381
‘Marina’ 43 mice 95, 252, 259, 558, 565, 576
Mark rootstock 95, 107, 109, 221, 256, 257 ‘Michinoku’ 40
marketing microbial insecticides 495
archetypes 563, 564 micropropagation 106, 127–128, 134
clubs 26–27 cultivar responses 136
consolidation 26 culture media 135
standards 410 ‘rejuvenation’ 136
Marssonina blotch 463 rootstock propagation 130–131
mass trapping 498 tree performance 49, 130
mating disruption 498 Mikado and Drilling systems 373, 382
maturity indices see harvest indices ‘Miki Life’ 40
maturity programmes 592–593 mineral
‘Maypole’ 37 elements
‘McIntosh’ annual requirements 267–268
bearing habit 74, 157, 160, 162, 328, 623 concentrations in fruit 268, 270–271, 569
breeding, use as a parent 33. 75 concentrations in leaves 268–270
chilling requirement 74 concentrations in roots 268
cold hardiness 63, 74, 75, 221, 222, 623 concentrations in tree framework 268
columnar types 75 critical values for plant growth 270
controlled atmosphere requirements 75, 596 nutrients, in soil 246–247
cultivar characteristics 74, 75 oils see oils
disease susceptibility 75, 561 mini central leader system
frost susceptibility 74 light distribution within canopies 388–390
fruit characteristics 74, 232, 413 planting densities 348
fruit growth 162 profitability 396–399
fruit quality 331 support system 380
global distribution 74 training method 348, 349–350
global production 18 use of semi-dwarfing rootstocks 348
impact of root pruning 336 yield comparisons 381
impact of summer pruning 333 mini-Tatura trellis system 372
insect pest susceptibility 75 mini-V-trellis system 373. 375
origin 74, 623 mirid pests 503
parentage 74 misting requirements of softwood cuttings 134
scald susceptibility 75 MM.106 rootstock
sports 43, 75 alternative rootstocks with similar vigour 103, 111,
spur types 43, 75 114–115
storage disorders 75, 607 collar rot susceptibility 95, 477
storage requirements 74–75, 596, 603, 607 propagation 133, 134, 135
tree form 74 yield efficiency 97
uses, as fresh and for processed products 74, 621, 623 MM.111 rootstock
see also 9, 12, 564, 588, 590, 592 drought tolerance 180
‘McIntosh Red’ see ‘McIntosh’ propagation 133, 135
‘McIntosh’ strains, responsiveness to controlled atmos- see also 257
pheres 586, 596, 601–602 MM series of rootstocks see Malling–Merton series
1-MCP see 1-methylcyclopropene models
‘McShay’ 64, 75 for evapotranspiration 182–183, 186
meadow orchard systems 363 for predicting apple scab infection 469–470
mealybugs 512, 546 for predicting chilling requirement 155, 239
mechanical for predicting fire blight infection 469
harvesting for predicting flower bud development 225
for processing grade fruit 627 for predicting powdery mildew infection 472
of Lincoln canopy 369, 371 for predicting yield 232
of Penn State trellis 368, 371 for water use estimation 186
650 Index
modified atmosphere (MA) storage 603–604 role in virus transmission 480, 481, 485
modified room cooling 597 suppression with cover crops 309
molecular maps 49 susceptibility of rootstocks 118
molecular markers 47–48 Neoseiulus californicus 510
see also 33, 37, 44 ‘New Red Star’ 17
‘Mollies Delicious’ 34 ‘New Spitzenberg’ see ‘Jonathan’
molybdenum 267, 270 nicotinoids (pesticides) 496
Monilinia leaf blight 463, 472 night temperature, effect on fruit set 428
Monochaetia twig canker (Seiridium unicorne) 463 nitidulid beetles 509
‘Monroe’ 72, 470, 624 nitrate 278–280, 309–310
mouldy core 463 nitrification 279
moult accelerating compounds (pesticides) 497 nitrogen
mowing equipment 315 annual requirement 281, 284
Mucor rot 68, 463 as ammonium 278
mulching as nitrate 278
as an alternative to herbicides 310 content in fruit 270–271, 281
impact on pests 310 content in leaves 246–247, 268–270, 281
impact on water retention 252 content in organic matter 570
in integrated production systems 542 content in tree framework 268
in organic production systems 252, 567, 568 content in trees 281
using specialized mowers 315 critical fruit concentration 270
with organic matter 252, 310, 567, 568 critical leaf concentration 270, 283
mullein plant bug (Campylomma verbasci) 64, 503 cycle 278, 279
multiple row planting systems 204, 206, 351, 353, 354, 386 deficiency 283
mummified fruit 474, 574 fertilizer use
‘Murray’ 75 in integrated production systems 542
mutation breeding 43 in organic production 277–278, 569–570
mutations, induced, natural and periclinal 43 in young plantings 395
‘Mutsu’ 39, 66, 157, 211, 221, 222, 328, 410, 470, 596, 621, flushing, use in CA storage 602
624 forms in soil 278, 280
mycorrhizae impact of pruning 322
impact on water uptake 173 impact on flowering 1, 286
impact on nutrient uptake 173, 286, 569 impact on fruit set 161
impact of weed competition 306
oversupply
NAA see naphthaleneacetic acid impact on environmental quality 280
naphthaleneacetic acid impact on fruit quality and storage disorders
application rates for control of preharvest drop 164, 283
443, 451, 452, 453, 454 impact on tree vigour 283, 523, 531
chemical formula 441 remobilization in trees 282–283
factors influencing effectiveness 227, 417, 426 role in metabolism 280
mode of action of action and use as a thinning agent soil availability 278, 279
411, 417, 418, 443 soil sampling methods 280
role in relation to ethylene production 337, 417, 418 supply
timing of application 427–428 as foliar sprays 285
use to control root suckers 445–446 leaching losses 279, 280, 284, 285
use to control water sprout growth 445 timing of applications 284
use to stimulate flower bud production 362, 363, 364, via fertigation 277, 285
429, 449 uptake 284
use with surfactants 421 see also 267
see also 439, 442 North Holland spindle system
‘Nabella’ 37, 562 planting densities 353
NAD see naphthalene acidimide production efficiency 204, 205
‘Nanna’ 41 pruning and training methods 353
‘Naoussa’ 38 use of dwarfing rootstocks 353
naphthalene acidimide (NAD) 417, 428, 422 ‘Northern Spy’
‘Natsumidori’ 39 cultivar characteristics 222
necrotic leaf blotch (of ‘Golden Delicious’) 476 precocity 154, 156
nectar 158 use as a rootstock 99, 118
Nectria canker (Nectria galligena) use in processing 620, 621, 623
control measures 477, 578 see also 157, 328
cultivar susceptibility 63, 66, 70, 72, 73, 75, 76, 79, 81 notching, use to stimulate branch formation 341, 445
impact of crotch angle 143 ‘Nova’ 39
rootstocks as hosts 118 ‘Nova Easygro’ 35
symptoms 463, 477 ‘Novamac’ 35
transmission during propagation 142 ‘Novaspy’ 35
Nectria twig blight (Nectria cinnabarina) 463 Novole rootstock 95
neem extract, use as a pesticide 492, 574, 576, 579 nurseries see tree nurseries
nematodes nursery tree quality
impact in replant sites 254, 485 importance of calliper 394
impact on nursery production 137 importance of ‘feathers’ 143, 258
Index 651
nutrient effect on ethylene production 600–601

exchange capacity 244 effect on respiration 600
management in organic production methods 277–278, in storage 596
555, 565, 568–570
uptake
impact of mycorrhizae 173, 286, 569 P.22 95, 103, 105, 130
impact of root growth 273 ‘Pacific Beauty’ 70
uptake mechanisms 273, 274 ‘Pacific Queen’ 70
nutritional value, vitamin C concentration 42, 47 ‘Pacific Rose’ 41, 70, 82, 476
paclobutrazol
impact on flowering 156
‘Obelisk’ 37 impact on fruit set 451
oblique palmette system 365–366 as an inhibitor of gibberellin biosynthesis 186, 439
oils palmette canopy systems
use in bloom delay 537 light interception 204
use as pesticides 497, 505, 579 performance characteristics 381–382, 396–399
use to substitute for chilling 239 ’regular’ and ‘free’ types 365–366
orchard floor management systems palmette-leader system
definition of 303 as a conversion method 330, 350, 390
role in environmental protection 304, 501 training method 251
orchard plant systems see planting systems pan evaporation, for estimating evapotranspiration 183
orchard site selection 238, 246, 293, 522–523, 538, 557–558 parasitic wasps 511, 513
organic parentage of specific cultivars 62, 64, 66, 68, 69, 71, 72, 74,
matter 75, 76, 77, 78, 80, 81
application rates 252 parthenocarpy 33, 48, 72, 81, 157
as soil amendments 252 partitioning 201–202
management in soils 304, 311 PBRs see plant bioregulants
nitrogen content 252 pelargonic acid 415
production Peniophora root canker (Peniophora sacrata) 464
adoption rate 554 Penman–Monteith equation 183
certification 554 Penn State thin-wall trellis system 368–369, 380
conversion requirements 555, 557 perennial canker (Neofabrae perennans) 464
cultivar selection 255, 560–562 periclinal mutations 43
ecological compensation 541, 599–560 permanent wilting point, definition of 245
fertilizer practices 277–278, 555, 565, 568–570 pest control
fruit thinning 556, 558–559 damage action threshold 305
grade standards 556–557, 581 economic injury level 305, 491, 499, 510, 540
market potential 554, 581 pesticide residues 490
mulches 252, 310, 567, 568 pesticide resistance 490, 544
pest management 499, 504, 556, 571–580 pesticides 492–498
planting stock management 555 petals 157, 158
planting systems 558 pH
postharvest management 556, 574 adjustment in soils 249–252, 291
principles 552–553 optimum values 247
pruning systems 556 phenological models for insect development 499–500, 507
returns 557, 581 pheromone traps
rootstock selection 559 for codling moth 576, 580
site requirements 557–558 for Graphania mutans 502
soil management 277–278, 555, 563–570 for tortricids 580
standards 553–554, 581 ‘Philip Rick’ see ‘Jonathan’
tree training 555, 558 Phomopsis canker, fruit decay and rough bark (Phomopsis
weed control 315 mali) 464
organochlorine pesticides 490 phosphate buffer capacity 286
organophosphate pesticides 490, 505, 507, 546, 548 phosphorus
organotins (pesticides) 494 application, in organic production 278, 570
oriental fruit moth (Grapholita spp.), feeding habit and availability in soils 246, 247, 286
control measures 506, 571 content in fruit 270–271
origin of Malus species 1, 92–93, 238 content in leaves 268–270
origin of specific cultivars 34, 40, 41, 62, 64, 66, 68, 69, 71, content in roots 268
73, 74, 76, 77, 78, 80, 328, 622, 623 content in tree framework 268
‘Orin’ 40, 51, 66 critical fruit concentrations 270
ornamental forms 35 critical leaf concentrations 270
osmotic adjustment in leaves 177, 178 fertilizer rates 287
osmotic potential foliar application method 277, 287–288
definition 168–169 impact of pruning 322
ovary 157, 158, 160, 161 role in low temperature breakdown 287, 288
overhead irrigation for frost protection 534–535, 538 role in metabolism 286–287
over-the-row mechanical aids 354 solubility in soils 276, 286
ovules 157, 158, 159, 160, 231 tree requirements 254
oxamyl, use as a thinning agent 417, 419 uptake by roots 172, 173, 286
oxygen concentration see also 267
652 Index
photomorphogenesis 197, 208, 209 pollination 157

photosynthesis polygenic resistance 45
canopy 180, 197–198, 199–200, 232 polyphenol oxidase 47, 51
effect of temperature 228–229 polyploidy 42
effect of water stress 178 pome, botanical description 10, 157
impact of cloudy weather 424–426 population growth models, of insects 499
impact of root pruning 334 postharvest
impact of summer pruning 198 cooling 597–598
inhibitors, use as thinning agents 418, 421 diseases 607–608
single leaf 197–199 see also bitter rot; black rot; blue mould; bull’s eye
photosynthetic photon flux, definition 196 rot; grey-mould rot; Penicillium spp.; white rot
photosynthetically active radiation (PAR), definition 196 disorders 604–607
Phymatotrichum root rot (Phymatotrychopsis omnivora) 464 see also bitter pit; brown core; carbon dioxide injury;
physiological disorders 593, 604–607 internal browning; low oxygen injury; low
physiological maturity 589–592 temperature breakdown; senescent break-
phytochrome 197, 208, 209 down; soft scald; superficial scald
Phytophthora crown, collar and root rot 464, 476 handling 585–586
Phytophthora spp. leaf retention, effect on productivity 232
cause of crown rot and root rot 477 management for organic production systems 556, 574
disease cycle and control measures 477 post-storage temperature control 598–600
interstem resistance 139 potassium
rootstock resistance 104–105, 108–110, 111, 112, availability in soils 246, 276, 288
114–115, 116, 118, 141, 477 content in fruit 270–271
see also 66, 464 content in leaves 268–270
phytoplasmas 478, 482 content in organic matter 570
picking platforms 365, 371 content in roots 268
‘Pilot’ 38 critical fruit concentrations 270
‘Pingo’ 38 critical leaf concentrations 270
‘Pink Lady®’ deficiency symptoms 288
global production 18, 64, 82 impact of pruning 322
marketing alliance 26–27 measurement method 288
origin 36 nitrate
parentage 66 role in bud break 225, 239
transformed selections 50 use as a foliar fertilizer 289
see also 470, 564, 596 relationship to calcium 288–289
pink mould rot (Trichothecium roseum) 464 relationship to magnesium 288, 289
‘Pinova’ see ‘Corail’ supply
‘Pioneer Mac’ 75 fertilizer forms and rates 289
‘Pionier’ 42 in organic production 278, 570
‘Pirella’ 38, 66 see also 267
‘Piros’ 38 powdery mildew (Podosphaera leucotricha)
PL system see palmette-leader system control measures 464, 471–472, 558, 573
plant bioregulants (PBRs) see growth regulators cultivar susceptibility 45, 68, 69, 70, 72, 73, 75, 76, 78,
plant protection in organic production systems 571–581 470, 573
planting epidemiology 470
density influence of humidity 471
for different systems 348, 353, 358, 362, 366, 369, 370, influence of temperature 471
372, 373, 375 molecular markers 47–48
impact on early yields 381–382 predictive model 472
impact on economic performance 377, 381, 382–384, qualitative and quantitative resistance 45
396, 398–400, 402 resistance, as a breeding objective, see breeding objec-
impact on fruit quality 330, 354 tives
impact on light interception 386–387 rootstock susceptibility 118, 128
influence on yield 204, 205, 206, 381, 382–384 symptoms 464, 470–471
stock quality see nursery tree quality precocity see rootstock selection
systems 346–402 precooling of fruit 597
comparisons of different types 380–382 predator:pest ratio 499
for organic production systems 558 predatory species for mite control 510–511, 575
time 259–260, 394 prediction of fruit size 162
ploidy predictive models
aneuploids, diploids and tetraploids 33 for apple scab infection 469–470
triploids 33, 72–73, 157 for fire blight infection 467
see also 11 for insect development 500, 507
plum curculio (Conotrachelus nenuphar) 502–503 for insect population growth 499
‘Pohorka’ 81 for light interception 202–204, 385
pollen 159 for powdery mildew infection 472
pollen tube growth 159–160, 227, 231 preharvest drop
pollenicides 415–416 control with AVG 451, 452
pollenizers 157–158 control with NAA 452, 453
planting patterns 158, 260 pressure potential 169
use of crab apples 159, 260 ‘Priam’ 66, 561
Index 653
‘Prima’ 49, 66, 561 of young trees 322, 324, 325–327

primary scaffolds 325 timing of 324–325
primary spur leaves 198–199 types of 322–323
‘Prime Red’ 39 Psylla mali 512
‘Primevere’ 35 pygmy fruit
‘Primicia’ 36 as a source of disease inoculum 423
‘Primiera’ 66 occurrence 418, 420, 421
‘Princesa’ 36, 66 pyramid canopy systems 206
‘Priscilla’ 64 pyramiding (of resistance genes) 45
‘Pristine’ 34 pyrethroid pesticides 490, 492, 495, 505, 507
process grade fruit standards 410, 620, 624–625, 627–631 pyrethrum, use as a pesticide 574, 575, 578
processed products
apple sauce 619–620, 621
baked apples 620 ‘Qiojin’ 37
cider 618–619 quantitative trait loci (QTL), for fruit texture 46
juice 616–618 quantum efficiency 198
sliced apples 620, 621 quassia, use as a pesticide 574, 575, 578
see also 19 ‘Queen Cox’ (‘Cox’s Orange Pippin’ sport) 81
processing apples quince rust (Gymnosporangium claviceps) 465
orchard management practices 625–627 ‘Quinguan’ 17, 37, 621
production volumes 616, 617
selection of cultivars 255, 621
production
radiation
cultivar trends 17
frost 240, 522, 531, 532, 534, 538
methods
interception, by earth 526
for 1-year-old trees 140–147
short-wave and long-wave components 524
for interstem trees 119–120
radiative freeze see radiation frost
world values 15–17
‘Ralls Janet’ 37, 39, 66
‘Produkta’ 37
randomly amplified polymorphic DNA 12, 44, 48, 49
prohexadione-Ca 442
RAPD see randomly amplified polymorphic DNA
as an inhibitor of gibberellin biosynthesis 439
rapid CA storage 597
chemical formula 441
reaction wood 337
role as an inhibitor of vegetative growth 186, 443
‘Reanda’ 38, 562, 564
use to promote fruit set 421
‘Rebella’ 38
see also Apogee®
receptacle tissues 158
Promalin®
rectangularity of plantings 260
application rates and timing 444–445
impact on productivity 377
impact on feathering 144, 145
influence on light interception 206
role in branching 444, 445
red:far-red ratio 197, 208
use to modify fruit shape 162, 453
‘Red Delicious’ see ‘Delicious’
see also 440, 441, 442
‘Red Earlilib’ 39
propagation methods for rootstocks 126–136
red skin colour
protection methods for freeze injury 220, 223–224
enhancement with ethephon 454–455
Provide®, application rates and timing to control russet
influence of solar radiation 163
and cracking 453
influence of temperature 232
pruning
‘Red Sparkle’ 35
definition 320
red spider mite (Panonychus ulmi) 63, 66, 69, 78, 476,
early records 320
510–511, 546, 575, 579
effect on branching 321
‘Redkroft’ 41, 222
effect on development of winter hardiness 321, 524,
reflectants, use to delay bloom 537–538
531
reflective mulches see light reflecting mulches
effect on flowering 322, 323, 428–429
‘Regali Delkistar’ 38
effect on mineral nutrients 322
‘Regent’ 34
effect on root growth 322
‘regular’ palmette system 365–366
effect on shoot growth 323
regulated deficit irrigation (RDI) 185–186, 187
effect on spur quality 322, 323
relative humidity
effect on yield 322
definition 529
for established trees 327
impact on freeze events 529–530
for processing apple trees 625
importance in fruit storage 600
impact on fruit set 322, 323, 428–429
role in energy balance 532–533
impact on light transmission 322, 323
relative water content, definition 169
impact on return bloom 428–429
‘Remo’ 38
impacts of heading cuts 323, 324, 392
‘Renetta Grigoa di Torriana’ 45
impacts of thinning cuts 323, 324
renewal pruning 355, 358, 391, 393
in relation to role of plant hormones 321
replant site management 100, 253
methods
‘Resista’ 37, 561, 564
for established trees 327
respiration rate
for old trees 330, 331
in controlled atmospheres 600
for organic production systems 556
of fruit 587, 589, 595, 600
for young trees 322, 324, 325–327, 334
respiratory climacteric 586, 595
of newly planted trees 261–262
654 Index
rest see dormancy for managing overcrowding 334, 336

restriction fragment length polymorphism (RFLP) timing of 335, 336
markers 49 restriction 97
ReTain® see aminoethoxyvinylglycine (AVG) rot see Phytophthora
return bloom suckers
impact of thinning on 414 control with NAA 446
promotion with ethephon 420, 429, 448 rootstock performance 102, 104–105, 109–110, 112,
promotion with NAA 362, 363, 364, 429, 449 114, 116–117, 257
role of gibberellins 439 susceptibility to fire blight 446
‘Rewena’ 38, 562, 564 use in propagation 7, 93
‘Rhode Island Greening’ 621, 623–624 root:shoot balance 320
‘Richelieu’ 35, 75 rooting depth 243, 272, 273
ring barking see ringing roots
ringing 339–341 cold hardiness 218, 242
impact on flowering 156 hydraulic resistance 169
impact on fruit quality 340 role in cytokinin biosynthesis 440
ripening 587 rootstock
date, heritability 44 breeding 99–100
‘Roanoke’ 77 selection
‘Rodluvan’ 42 fire blight susceptibility 95, 107–108, 111, 112,
Roman Empire, spread of cultivation 7, 93 114–117, 118
‘Rome Beauty’ for different orchard systems 384–385
bearing habit 76, 157, 328, 622 for orchard establishment 196
breeding, use as a parent 77 for organic production systems 100, 119, 559
chilling requirement 76 for vigour control 102–117
cold hardiness 76 graft compatibility 101
controlled atmosphere requirements 76 health status 101
cultivar characteristics 76 impact of site 106
disease susceptibility 76, 470, 475 impact on fruit size 104, 108–110, 112, 114, 257, 384
frost susceptibility 76 influence on precocity 94, 96, 97, 101, 104–105, 106,
fruit characteristics 76, 622 108–110, 111, 112–113, 114–115, 116–117, 256,
global distribution 76 384
global production 18 influence on yield 96, 256
insect pest susceptibility 66, 76 influence on yield efficiency 104–105, 106, 108–110,
origin 76, 622 112–113, 114–115, 116–117, 256–257, 384
parentage 76 resistance to woolly apple aphid 95, 100, 111,
pollination requirement 157 114–115, 116, 118, 257, 384
pruning options 325 site index 96
scald susceptibility 76 transformation 51, 100
sports 77 rootstocks
storage requirements 76 AR series 103
tree form 76, 622–623 collar/crown rot resistance 104–105, 108–110, 111,
uses, as fresh and for processed products 76, 620, 621, 112, 114–115, 116, 118
623 description 92
see also 77, 588 dwarfing selections 103, 106–107, 108–110
‘Romus 1’ , ‘Romus 2’ and ‘Romus 3’ 42 ease of propagation 101, 103, 104, 106
root fire blight resistance 108, 111, 112, 114–115, 116–117,
density 170, 171, 181, 272, 303–304 118, 257, 384
distribution history of use 92, 94
impact of root pruning 336 impact of environment 95, 96
impacts of irrigation emitters 177, 273, 276 impact on fruit maturation 94, 257
growth impact on fruit set 94, 96
effect of soil water potential 180 impact on growth habit 94, 96, 98
effect of thinning time 414 impact on productivity 98, 384–385
impact of crop load 173, 181 impact on terminal bud formation 98
impact of pruning 322 impacts on orchard performance 106, 384–385
impact on nutrient uptake 273 importance in orchard systems 106
impact on water uptake 179 importance of disease resistance 95, 101, 118
periods of active growth 173, 305 importance of pest resistance 118
response to temperature 230–231 influence on drought tolerance 93, 95, 98, 107, 119,
induction, role of IBA 442 127, 384
initiation, role of IAA 439 influence on vigour 101, 255–256, 347
pruning 72 interactions with scions 95
effect on cytokinin synthesis 334 M.26 51, 94, 95, 99, 100, 102, 103, 107–108, 112–113,
effect on ethylene production 334, 336 130, 133, 134, 135, 180
effect on leaf water potential 334 M.27 94, 95, 96, 99, 103, 104–105, 128, 133, 180, 204,
effect on nursery trees 336–337 205, 256, 257
effect on root growth 335, 336 M.7 51, 95, 99, 100, 111, 257
effect on vegetative growth 97, 335, 337 M.9 93, 94, 95, 96, 99, 100, 103, 106–107, 108–110, 128,
effects on fruit quality, fruit size and yield 336, 337 129, 130, 133, 135, 137, 141, 180, 204, 205, 222,
effects on photosynthesis and transpiration 334 257, 258, 351, 385
Index 655
Malling series 99, 347 seaweed, use as a fertilizer 570

Malling–Merton series 99, 118 seed
Mark 95, 107 colour, as a harvest index 589
Merton 793 99, 118 development 160
Merton Immune series 99 role in fruit set 413
micropropagation methods 106, 135–136 number 157
MM.106 95, 97, 111 impact on fruit size 413
mode of action 96–97, 98 orchards 126
Northern Spy 99, 118 propagation 126
nursery performance 101 stratification 126–127
propagation 106 seedlings, use as rootstocks 92, 93, 102, 126, 347
semi-dwarfing selections 103, 107, 111, 112–113 seeds 158
semi-vigorous selections 103, 111 source of gibberellins 155
suckering habit 101, 257 ‘Sekaiichi’ 39, 64, 66
super dwarfing selections 103, 104–105 ‘Selena’ 37
tolerance of winter injury 92, 95, 99, 101, 104–105, self incompatibility 32, 33, 48, 157
107, 108–110, 111, 112–117, 118, 221, 222, 384 semi-dwarfing selections, of rootstocks 103, 107, 111,
use of seedlings 92, 93, 102, 126 112–113
very vigorous selections 111, 116–117 semi-hardwood cuttings 127
vigorous selections 111 semi-vigorous selections, of rootstocks 103, 111
virus status 101 senescent breakdown 72
Rosellinia root rot (Rosellinia necatrix) 464 control measures 605–606
‘Rosenapfel’ 45 ‘Senshu’ 40, 68
‘Rosu de Cluj’ 42 sensory testing 46
rosy apple aphid (Dysaphis devecta, Dysaphis plantaginea) ‘Seokwang’ 40
64, 66, 75, 513, 559, 574, 579 sepals 157
rosy leaf-curling aphid 48, 574, 579 ‘September Ruby’ 35
Rotenone , use as a pesticide 478, 492, 575, 578 ‘Septer’ 72
‘Rouville’ 35 shade
row orientation impact on fruit abscission 160–161, 208, 209, 424–426,
effect on productivity 202–203, 329 427
impact on orchard layout 258 impacts on flower bud differentiation 198–199, 208
‘Royal Gala’ see ‘Gala’ impacts on fruit mineral content 208
‘Rubens’ 39 impacts on fruit quality 208, 230
‘Rubinovoe Duki’ 72 impacts on fruit set 201, 208
russet impacts on fruit size 201
control using gibberellins 79, 81, 417, 453 shade leaves, characteristics 174, 198, 208
on fruit 226 SHAFFE see Southern Hemisphere Association of Fresh
russeting, heritability 44 Fruit Exporters
‘Russian seedling R12740–7A’ 44 ‘Shamrock’ 35, 66
rusts 464 ‘Shizuka’ 66
Ryania extract, use as a pesticide 492 shortwave radiation 196
side grafting 143
side rot (Phialaphora malorum) 465
salinity 247–248 ‘Silken’ 35
‘Sampion’ 81 silver leaf (Chondrosterium purpureum) 66, 70, 465
San Jose scale (Quadraspidotus perniciosus) 512 simazine 248
‘Sansa’ 40, 70 simple sequence repeats (SSRs), use in cultivar identifica-
‘Santana CPRO’ 41, 79, 562, 564 tion 48
SARD see specific apple replant disease ‘Sinsekai’ 40
‘Saturn’ 37, 561 ‘Sir Prize’ 66, 561
‘Sawa’ 41, 222 site
sawflies see apple sawfly index (for rootstock selection) 96
scab see apple scab preparation 248, 249
scald see superficial scald selection
scale insects 491 for freeze avoidance 522–523, 538
scarab beetles 514 for highly productive orchards 246, 293
scarf skin 76 for organic production 557–558
‘Scarlet’ 40 ‘Siv’ 41
‘Scarlet O’Hara’ 34 ‘Skala’ 42
‘Scarlett Sentinel’ 35, 75 skeletonising insects 511
‘Sci Early’ 41 skin, resistance to gaseous diffusion 601–602, 603
‘Sci Red’ 41 skin temperature 230
scion inclination, effect on productivity 339 slender-pyramid system
scions, transformation 50, 51 planting densities 357
scolytid beetles 514 pruning method 358, 359
scoring 339–341 support system 380
to control vigour 340, 448 training method 357–358, 359, 392
impact on flowering 156 use of semi-vigorous or dwarfing rootstocks 357
impacts on fruit quality and fruit size 340 slender-spindle system 365
see also 355, 359 gas exchange characteristics 200, 202, 203
656 Index
slender-spindle system continued see also Malus species

light distribution within canopies 388–390 specific apple replant disease (SARD) 137, 253, 309, 395
light interception 386–387 spectral distribution, influence of leaf canopy 196–197
planting densities 351, 353 ‘Spencer Seedless’ 44, 155
profitability 396–400, 401, 402 spherical shaped canopies 347–348
support system 380 bush-tree system 344
training method 207, 330, 339, 351, 352, 353 spider mites (Panonychus ulmi, Tetranychus spp.) 490,
use of dwarfing rootstocks 351, 354 510–511, 575
yield efficiency 204, 205, 206 spindle-bush system 347
yields 256, 381, 382 ‘Splendour’ 41
sliced apples sports of specific cultivars
grade standards 624, 625 ’Braeburn’ 78
processing methods 620 ‘Cox’s Orange Pippin’ 81
‘snip’ trees, production methods 138–139, 140, 146 ‘Delicious’ 43, 64
soap (use as a pesticide) 498, 574, 575, 579 ‘Elstar’ 79–80
soft scald 597, 605 ‘Empire’ 43
symptoms and control measures 606 ‘Fuji’ 43, 68
see also 597, 605 ’Gala’ 43, 70–71
softwood cuttings 127, 133–135 ‘Golden Delicious’ 43, 66
soil ‘Granny Smith’ 69, 97
aeration 244, 523 ‘Jonagold’ 43, 73
characteristics, in relation to site selection 394, 558 ‘Jonathan’ 72
compaction 243, 523 ‘McIntosh’ 43, 75
contaminants 248 ’Rome Beauty’ 77
cultivation 248 spotted cutworm (Xestia c-nigrum) 509
depth 243 spreaders 156, 338, 339, 349
drainage 245, 249 spur
fertility, management of 304 bearing habit 328, 522
fumigation 253. 254 leaf area, impact on fruit size 162
in relation to specific apple replant disorder 253, 395 leaves 201
management, for organic production systems pruning 330
564–565 quality
moisture effect of thinning time 414
measurement 182 impact of pruning 322, 323
potential, estimation 169–170 impact of root pruning 335, 337
organic matter 252, 304, 558
impact of summer pruning 331
pH see pH
types
quality, for tree raising 136–137
of ‘Delicious’ 64, 328
structure 243, 558
of ‘McIntosh’ 43, 75, 328
testing, for nutrient status 273–274, 280, 286, 288, 289,
spurs 157
293, 295, 296, 297, 298
Spy decline see apple stem pitting
texture 242–243, 244, 245
SRD see specific replant disease
water holding capacity 244, 245, 523
SSG see sylleptic shoot growth
solar constant 196
stamens 157, 158
SolAxe system, pruning and training methods 357, 392
Solen system ‘standard’ trees 94, 558
pruning method 370 standard CA storage 602–603
training method 357, 370 starch concentration 589
use of dwarfing rootstocks 370 starch index 589, 590, 591, 592
yield comparisons 382 ‘Stayman’ 453, 624
soluble solids concentration, as a maturity index 589, 590, Steiner, Rudolf 553
592 ‘Stellar’ 34
solute potential stem elongation, role of gibberellins 439
adjustment of 179 stem water potential 171, 181
definition 168–169 sterol inhibitors, use as thinning agents 421
somaclonal variation 49 stigma 158, 159
sooty blotch complex 465 stink bugs (Acrosternum hilare and Euschistus conspersus)
causal organisms 474–475 509
control measures 475, 573, 578 stomatal conductance
symptoms on fruit 474 control of water use 170, 183
source: sink relationships, influence of light 201 impact of crop load 175
South American fruit fly (Anastrepha fraterculus) 507 impact on evapotranspiration 171, 183
southern blight (Sclerotinium rolfsii) 465 modelling of 186
Southern Hemisphere Association of Fresh Fruit stool beds see stooling
Exporters (SHAFFE) 28 stooling 128–129
‘Southern Snap’ 41 see also 94
‘Spartan’ 35, 73, 75, 221, 222, 328, 561, 564, 592 storage
species of fruit
distributions 11 disorders see postharvest disorders; postharvest dis-
sections 11 eases
use as ornamentals 2–6 effects of high carbon dioxide concentrations 600,
use as rootstocks 2–6 607
Index 657
effects of low oxygen concentrations 600 control measures 593, 604, 606
effects of relative humidity 600 cultivar susceptibility 63, 65, 67, 69, 70, 72, 75, 76, 77,
effects of temperature 596, 598, 604 79, 80, 593, 596
life effect of water stress 178
impact of ethephon 454 impact of temperature 232, 605
impact of harvest period 587–588, 592–593 use of DPA 593, 603, 606
quality, effect of summer pruning 331 superior oil, use with thinning agents 421
requirements of specific cultivars 63, 65, 67, 69, 70, 71, support requirements 257
73, 74–75, 76, 77–78, 79, 80, 596, 603, 607 support systems see tree support
scald see superficial scald ‘Sweet Caroline’ 66
temperature 596, 598 ‘Sweet Sixteen’ 34
Streif Index 589, 592 sylleptic shoot growth 231
string tree system 362 sylleptic shoots 143
stub pruning 361 ‘Sylvia’ 42
styles 157, 158
sublimation (of water) 529
suckers see root suckers; water sprouts tabletop bed systems 371
sulphur ‘Takane’ 40
availability in soils 246, 292 tarnished plant bug 503, 579
content in fruit 270–271 taste
content in leaves 268–270 as a breeding objective 46
content in organic matter 293 impact of aminoethoxyvinylglycine 455
content in roots 268 Tatura trellis system
content in tree framework 268 planting density 372
critical leaf concentrations 270 training method 339, 371, 372
deficiency symptoms 293 use of semi vigorous rootstocks 372
role in metabolism 292–293 yield comparisons 381
sources of 293 taxonomy 9–11
use as a pesticide 492 T-budding 141–142
see also 267 ‘Telamon’ 37, 75
summer pruning temperature
effect on flowering 333 change with elevation 238, 239
effect on fruit colour 72, 331, 332, 333, 334 effect on cell division 161
effect on fruit quality 322, 331, 332, 333 effect on ethylene production 595, 597
effect on fruit set 331, 333 effect on flower production 230
effect on fruit size 333 effect on fruit growth 161, 229–230
effect on photosynthesis 331, 333 effect on fruit maturation 229–230
effect on spur quality 331 effect on fruit metabolism 595–600
effect on tree water use 174 effect on fruit quality 230, 232
effect on vegetative growth 333, 334 effect on fruit respiration 595, 597
effect on within canopy light distribution 207, 332, 390 effect on harvest date 228
in multiple-row systems 354 effect on insect growth 500
timing and severity of 331, 333–334 effects on leaf assimilation and respiration rates
‘Summer Treat’ 34 228–229, 232
‘Summerdel’ 64 effects on shoot and root growth 230–231
‘Summerred’ 35, 66, 75, 120, 228, 561, 564 impact of continental climate 218, 238
sun leaves, characteristics 198, 208 impact of maritime climate 218
sunburn on fruit impact on apple scab infection 469, 470
control methods 230 impact on crop yield 231, 232
control using tree training 230, 258, 360, 378 impact on feathering 231
cultivar differences 64, 66, 68, 75, 230 impact on fire blight infection 467
effectiveness of reflective coatings 212, 230 impact on fruit storage potential 232
impact of shade 212, 230 impact on fruit set and fruit thinning 227, 424–426,
impact of temperature 212, 230 428
symptoms in storage 605 impact on pollen tube growth 227
‘Suncrisp’ 34, 81 impact on powdery mildew infection 471
‘Sundowner’ 36, 66 impact on red skin colour 211, 232
‘Sunrise’ 35, 221, 222 influence on rooting cuttings 132–133, 134
sunscald see sunburn on fruit inversion 226, 240, 526–528
super dwarfing selections, of rootstocks 103, 104–105 monitoring 531–532
super-spindle systems requirements for fruit storage 596, 598
mineral nutrient requirements 268 ‘Tentation’ see ‘Delblush’
orchard life 363 tephritid fruit flies 506–507
planting densities 362 terbacil 248
profitability 400, 401 terminal buds, importance in flowering 157
pruning and training methods 362, 363, 364 tetranychid mites 490, 510–511, 575
support system 380 see also European red mite; red spider mite; spider
use of plant growth regulators 362, 363 mites
yield comparisons 381 texture
supercooling 222–223 as a breeding objective 46
superficial scald heritability 44
658 Index
Thinex 415 rootstock spacing 137

thinning shoot removal 139–140
agents site selection 136–137
Accel® 417, 420, 440, 441, 442 tree defoliation 146–147
ammonium thiosulphate 227, 415 tree lifting 147
application rates 422 tree spacing 137
benzyladenine 411, 417, 420–421, 441 tree support 139
carbaryl 411, 417, 418–419, 420, 421, 428 see also budding height; budding methods; bud
DNOC 413, 415 wood; grafting
Dylox 417, 420 planting depth 260
effectiveness 423, 426, 428 planting time 259–260, 394
Elgetol 413, 415 quality 258–259
Endothall 415 importance of calliper 394
ethephon 417, 419–420, 428, 448 importance of ‘feathers’ 139, 394
herbicides 421 stakes 379
long-chain fatty acids 415 storage 147, 259
morestan 417 support
NAA 411, 417–418, 443 costs 261, 378, 380, 396
NAD 417, 418, 422 in the nursery 139
oxamyl 417, 419 in organic production systems 559
pelargonic acid 415 in relation to rootstock selection 260
photosynthesis inhibitors 418, 421 systems 261, 378–380
sterol inhibitors 421 training
Thinex 415 definition 320
Thinset 415 methods 347–377
Wilthin 415 systems, for organic production 555, 558
YI-1066 415 vigour
cuts 322–324 response to pruning 320–322
time vegetative:fruiting balance 392–393, 444,
impact on fruit size 411, 413, 414 446
impact on vegetative growth 414 tree water status, measurement of 170
see also fruit thinning ®
Tree-Hold 442, 445, 446
Thinset 415 tree-row-volume (TRV) spray rates 422, 443–444
thiourea 239 tri-iodobenzoic acid, impact on flowering 156
thread blight (Corticium stevensii) 465 triploid cultivars 33, 72–73, 157
thrips, feeding habit and damage symptoms 503 trunk injury
tip bearing habit 328–329
from cultivation 565
Tipoff® 446
from freeze damage 241, 242, 531
tipping, for control of feathering 143–144
TRV see tree-row-volume
titratable acidity use as a harvest index 589, 590, 592
‘Tsugaru’ 39, 66
tobacco budworm 576, 579
turgidity, of tissues 169
tomato ringspot nepovirus (TmRSV)
turgor potential
diagnostic methods 481
definition 168, 169
rootstock sensitivity 99
maintenance of 179
transmission vectors 481
‘Tuscan’ 37, 75
top grafting 140
‘Twenty-Ounce’ 624
top working see top grafting
‘Topaz’ 82, 562, 564 twospotted spider mite 64, 66, 68, 489, 510, 575
top-working disease 480 ‘Tydeman’s Red’ 75, 325, 328, 592
tortrix moth 576, 579, 580 tying methods, use in budding 142
trafficability, of soils 243, 244 Typhlodromus occidentalis 510
‘Trajan’ 37, 75 Typhlodromus pyri 510, 546, 579
transformation, of rootstocks and scions 50, 51
transgenic
cultivars 43, 46, 51 ‘Udine’ 72
rootstocks 46, 51, 100, 107–108 ULO see ultra low oxygen
transpiration rate 169, 186 ‘Ulster’ see ‘Jonathan’
transport of fruit 599 ultra low oxygen (ULO), in relation to CA storage 65, 67,
tree 69, 71, 79, 80
arrangement in nurseries 137 ultra low volume spraying, for application of thinning
form of specific cultivars 62, 65, 67, 68, 70, 71, 72–73, agents 422
74, 76, 77, 78, 80, 328, 329, 330 ultraviolet radiation (UV), influence on fruit colour 208,
form types 328–329 211, 212
height under tree irrigation for frost protection 535, 538
in relation to alley width 204, 329, 387 uniconizol, as an inhibitor of gibberellin biosynthesis
in relation to temperature inversion 523 439
lifting in nurseries 147 urea, use as a foliar fertilizer 161, 277, 285–286
nurseries uses of specific cultivars, as fresh and for processed prod-
’bleeding syndrome’ 138 ucts 63, 65, 67, 69, 70, 71, 73, 74, 76, 77, 79, 80, 620,
’feathering’ 138, 143–145, 258 621, 622, 623, 624
heading 137–138 Utah model, for chill unit determination 155
Index 659
V slender-spindle system see Gütingen V slender-spindle WAPA see World Apple and Pear Association
system water
V super-spindle system 375 loss, from fruit 585
Valsa canker (Valsa ceratosperma) 63, 68, 465 potential 168, 179
Vapam 254 sprinkling
variegated cutworm (Periodroma saucia) 509 for bloom delay 226, 537
vegetative growth for frost protection 226, 534–535, 538
control with Apogee® 443 for prevention of sunburn 230
effect of thinning time 414 sprouts
impact of branch bending 337–338 control with NAA 445
impact of root pruning 97, 335, 337 removal by summer pruning 334
impact of summer pruning 333, 334 stress 176, 177–179
impact of water stress 176 uptake
vegetative:reproductive balance 392–393, 444, 446 impact of mycorrhizae 173
Venturia inaequalis see apple scab impact of root growth 179
vertical-axis system use
fertilizer management 395 coefficient
planting densities 354 impact of humidity 184
profitability 396–399 impact of orchard layout 183–184
pruning method 354, 355–356 crop coefficient 183–184
support system 380 efficiency (WUE) 170, 179, 203
training method 354, 355–356, 392 impact of canopy form 175
use of dwarfing rootstocks 354 impact of drought 175
yield comparisons 381–382 impact of environment 173, 181–184
yields 256 impact of leaf area 174, 175
see also 365 impact of solar radiation 173–174
very vigorous selections of rootstocks 111, 116–117 impact of summer pruning 174
vesicular arbuscular mycorrhizae (VAM) 173, 254, impact of tree spacing 175
286 rates of 173
vespid wasps 509 water-core 67, 72, 232, 455, 589, 604
vigorous selections of rootstocks 103, 111 waxes, as fruit coatings 594
vigour ‘Wayne’ 621
control weed management
impact of pruning 322–324, 327 control thresholds 305
influence of rootstocks 94 impacts of irrigation 306
selection of rootstock 102–117 in organic systems 563–564
use of interstems 92, 119 in young plantings 172, 306, 394
heritability 44 weeds
types (of cultivars), classification 327–329 control using heat 315, 567–568
violet root rot (Helicobasidium mompa) 465 control using herbicides 305, 307, 309, 313, 314
viroids 478, 482–483 effect on rootstocks 559
virus hosts of specific viruses 312
diseases 478–483 impact on winter injury 523
virus infection, effect on rootstock propagation 130, impacts on productivity 304, 394
140 weevils 502–503
viruses 140, 478, 485 ‘Wellington Bloomless’ 44
virus-like diseases 478, 479, 483–485 western box-elder bug (Leptocoris rubrolineatus) 509
visible light 196 western flower thrips (Frankliniella occidentalis) 490, 503
‘Vista Bella’ 34, 222, 561, 564 western yellowstriped armyworm (Spodoptera praefica)
vitamin C concentration, selection in breeding pro- 509
grammes 42, 47 wettable sulphur 577–578
‘Voinea’ 42 whip and tongue graft 143
voles 95, 252, 259, 310, 558, 565, 576 whips, pruning management 325, 351, 352, 360
V-shaped canopies ‘White Angel’ 48
light distribution within canopies 388, 389, 391 white root rot (Scytinostroma galactinum) 465
light interception 377–378, 386–387 white rot (Botryosphaeria dothidea)
performance characteristics 377–378, 381–382, 387 control measures 465, 474, 607
profitability 396–399 symptoms on fruit 465, 473, 474
V-shaped canopy types 371–378 whole canopy gas exchange
Geneva Y-trellis system 372–373, 374, 380, 396–399 impact of canopy form 200, 203–204
Gütingen V slender-spindle system 375, 376–377, impact of temperature 228–229
396–399 measurement 199
MIA trellis system 373, 381 relationship to yield 200
Mikado and Drilling system 373, 382 relationship to light interception 203
mini-Tatura trellis system 372 ‘Wijcik McIntosh’ (‘Starkspur Compact Mac’) 43, 49, 75,
mini-V-trellis system 373, 375 97
Tatura trellis system 339, 371, 372, 381 Wilthin 415
V super-spindle system 375 wind machines
V slender-spindle system see Gütingen V slender-spindle for frost protection 535–536, 538
system mode of action 527, 528, 536
V super-spindle system 375 use with heaters 536
660 Index
‘Winesap’ 62, 157, 410, 620, 621 efficiency, impact of rootstock 104–105, 106, 108–110,
‘Winston’ 81 112–113, 114–115, 116–117, 256–257, 384
winter impact of fruit thinning 413
chilling 155 influence of rootstock 96
dormancy see dormancy relation to light interception 195, 197, 202, 385–387
hardiness relation to temperature 231
impact of pruning 321, 524, 531 yields, in organic production systems 581
influence of rootstocks 111 ‘Yoko’ 40
see also freeze injury ‘York Imperial’
moth 576, 579 cultivar characteristics 622, 625
‘Witos’ 41 use in processing 621
wood boring insects 513–514 see also 157, 410, 417, 420
woolly apple aphid (Eriosoma lanigerum) young tree establishment
biological control 513, 546, 574 competition from weeds 559
resistance role of irrigation 394
markers for resistance 48 young trees
of rootstocks 95, 99, 100, 111, 114–115, 116, 118, 257, defoliation 146
513 pruning strategies 325–327, 334
see also 64, 69, 514 Y-trellis 204, 206, 207, 372, 373, 374, 381–382, 395, 396–399
World Apple and Pear Association 28 see also V-shaped canopies; V-shaped canopy types
‘Xinguan’ 66 zeatin 439, 440

‘XinShuai’ 37 ‘Zestar’ 34
Xiphinema americanum 480, 481, 485 zinc 267
xylem, low temperature threshold 218, 220, 223 availability in soils 246, 247, 295
critical leaf concentrations 270
deficiency symptoms 295–296
‘Yanshanhong’ 37 role in metabolism 295
‘Yellow Newtown’ 157 supply as foliar sprays 277, 296
YI-1066 415 ‘Zoete Oranje’ 81
yield zonate leaf spot (Crustulariella moricola) 465
effect of pruning 322 ‘Zuzana’ 37, 562

Apples

Uploaded by

Apples

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1.1.

Plate 3.5. Colour sports of ‘Gala’ developed naturally as mutations.

Plate 4.5. ‘Imperial Gala’ (from Bruce Barritt).

Plate 6.4. Mature layer bed of M.9.

Plate 6.14. Tying the chip to the rootstock.

Plate 11.3. Orchard in British Columbia, Canada, planted across a hillside.

Plate 12.8. Zinc-deficiency symptoms manifested on young ‘Delicious’/M.9 apple trees.

Plate 14.3. Mechanical root pruning of apple trees at bloom.

Plate 15.12. Geneva Y-trellis apple orchard of ‘Empire’ on M.26 rootstock.

Plate 20.3. Towers used to monitor inversions and wind-machine effects.

Plate 23.1. Fruit inspection in a modern packing-house.

Botany, Production and Uses

The editors dedicate this book to all the professors,

Botany, Production and Uses

Department of Horticulture and Crop Science, Ohio State University, USA

Department of Horticultural Science, Massey University, Palmerston North,

CABI Publishing is a division of CAB International

Tel: +44 (0)1491 832111 Tel: +1 617 395 4056

© CAB International 2003. All rights reserved. No part of this publication

Library of Congress Cataloging-in-Publication Data

ISBN 0 85199 592 6

Typeset in 9 on 11 pt Palatino by Columns Design Ltd, Reading

PART II: PLANT MATERIALS

PART III: APPLE PHYSIOLOGY AND ENVIRONMENTAL INFLUENCES

PART IV: ORCHARD AND TREE MANAGEMENT

13. Orchard-floor Management Systems 303

PART V: CROP PROTECTION

PART VI: HARVESTING, HANDLING AND UTILIZATION

Colour plate section after p. 436

1 Taxonomic Classification and Brief History

1.1 The Origin and Spread of the Domesticated Apple 1

© CAB International 2003. Apples: Botany, Production and Uses

Taxonomic Classification and History

M. prunifolia (Willd.) Borkh. 34 Central and eastern China Ornamental, rootstock

Taxonomic Classification and History

M. × robusta (Carriere) [= M. baccata × M. prunifolia]

Taxonomic Classification and History 7

revered in keeping with Koranic teachings Apples were introduced to Australia, on

Taxonomic Classification and History 9

nations. Later in the century, the USSR also

1.2.1 Apples in the family Rosaceae

Subfamily Subfamily Subfamily Subfamily

Genus Genus Genus 20 other

Section Section Section Section

Series Series Series Series Series Series

Fig. 1.2. Taxonomy of Malus (adapted from Phipps et al., 1991).

Taxonomic Classification and History 11

Taxonomic Classification and History 13

2 World Production, Trade, Consumption

2.1 Introduction and ‘Fuji’) and advances in irrigation tech-

© CAB International 2003. Apples: Botany, Production and Uses

Apples: World Production, Trade and Consumption 17

they could receive. A smallholding that 2.3 Commercial Practices

2.4 Cultivars the apple business at the expense of existing

Table 2.1. Major producing countries excluding China, cultivar trends

2000 2005 2010

‘Red Delicious’ 5,334 5,422 5,423

Grand total 26,313 29,131 30,429

Apples: World Production, Trade and Consumption 19

Country/region 1993/94 1994/95 1995/96 1996/97 1997/98 1998/99 1999/2000p

N. hemisphere 509,306 509,999 454,797 451,090 519,852 515,128 500,500

World total 654,360 651,804 611,837 640,504 672,095 678,402 632,662

N. hemisphere 200,531 208,446 107,457 90,241 132,520 129,834 97,495

World total 201,206 209,331 107,957 97,589 140,685 132,929 99,090

Apples: World Production, Trade and Consumption 21

1 France 766,207 488,559 Germany 707,763 438,656

Top ten 4,089,791 2,233,314 Top ten 2,686,379 1,764,234

Apples: World Production, Trade and Consumption 23

n/a, not available; p, preliminary; EU, European Union.

Apples: World Production, Trade and Consumption 25

Country and region 1990 1994 1995 1996 1997 1998

needed to achieve a given percentage most systematic approach to studying these

Apples: World Production, Trade and Consumption 27

Apples: World Production, Trade and Consumption 29

3 Genetic Improvement of Apple: Breeding,

Susan K. Brown and Kevin E. Maloney