Principles of Grafting
Principles of Grafting
Principles of Grafting
11
Principles of Grafting
and Budding
INTRODUCTION learning objectives
• Describe the role of grafting in
Since the beginning of civilization, fruit and nut trees have been grafted
human history.
because of the difficulty in propagating by cuttings, and the superiority
• Distinguish between the
and high value of the grafted crop. Grafting is among the most expensive
use of seedling and clonal
propagation techniques, surpassing even micropropagation. Budding,
rootstocks.
which is a form of grafting, is three times more costly than cuttings and
• Describe how natural grafting
fourteen times more expensive than seedling propagation (89). The horti-
can affect tree performance.
culture and forestry industries have sought to develop clonal propagation
• Describe how the rootstock
systems that avoid labor-intensive graftage. Yet, traditional and highly
and scion heal together during
efficient grafting and budding systems are essential for the propagation of grafting.
many woody plant species. New markets continue to require grafted and
• Define how specific genetic,
budded plants for improved plant quality, fruit yield, superior forms, and environmental, and manage-
better adaptation to greater ecological ranges. In the southeastern United ment factors and polarity
States, where high temperatures and periodic flooding of soils (low soil affect graft success.
oxygen) are the norm, cultivars of birch, fir, oak, and other species are • Determine what kinds of
grafted onto adapted rootstock (Fig. 11–1) (129). The propagator bene- plants can be grafted.
fits via new markets, while the consumer gains a greater variety of better- • Define graft incompatibility—
adapted landscape plants. The acid-loving blueberry can be produced in its symptoms, causes, and
more basic pH soils when grafted to pH-tolerant rootstock (Fig. 11–2). control.
With the greater reliance on integrated pest management and • Describe important ways the
reduced availability of pesticides and soil fumigants, disease-tolerant rootstock (root system) influ-
rootstocks are playing a greater role not only with woody perennial fruit ences the scion (shoot system)
crops and ornamentals, but also with grafted vegetable crops (Figs. 11–3 and vice versa.
and 11–4, page 417) (34, 39, 67, 82, 85, 86). Organic growers of high
value heirloom tomatoes are using grafted plants as a management tool
to reduce crop loss from soilborne diseases (131).
This chapter reviews the biology of grafting and budding.
Chapters 12 and 13 describe the techniques of grafting and budding,
respectively. Chapter 19 enumerates grafting and budding systems for
selected fruit and nut trees, as Chapter 20 does for selected woody orna-
mental plants. A better understanding of the fundamental biology of graft-
ing (and the causes of graft incompatibility) will enhance the development
of superior cultivars and increase the ecological range of species for new
markets in horticulture and forestry.
Figure 11–1
Cleft-grafted-variegated
English Holly on Ilex ‘Nellie
Stevens’ rootstock adapted
to the high temperature,
periodic flooding, low oxygen
soils of the southeastern
(a) (b) United States.
Figure 11–2
Pushing the ecological
envelope. Using an inlay bark
graft of ‘Tif Blue’ blueberry
(Vaccinium ashei) on a
farkelberry (Vaccinium
arboreum) rootstock, which
tolerates a more basic soil
pH, allows the acid-loving
blueberry to be produced
in a site with higher soil pH.
(a) New scion growth with
aluminum foil and poly bag
protecting the graft area.
(b) Healed graft union, and
(a) (b) (c) (c) ‘Tif Blue’ blueberry crop.
Scion
Non-Grafted
Grafted
Rootstock
(a)
(b)
Figure 11–3
Grafting vegetables is a common practice in Japan, Korea, the Mediterranean basin, and Europe. It is used for managing soil-borne
diseases, enhancing tolerance of low temperature and salinity, and for increasing plant vigor and yield. (a) Grafted melon scion
on curcurbita rootstock with a grafting clip. (b) Melons grafted (white arrow) on Fusarium-resistant Curcurbita rootstock in Israel,
(b) compared to susceptible, non-grafted melons (black arrows). Courtesy M. Edelstein.
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Early crop
(a) (b)
Grafted Non-grafted
Figure 11–4
(a and c) Melon grafted onto
boron-resistant Cucurbita
rootstock. (b) Non-grafted
melon showing boron
susceptibility early in crop
1-month later
cycle and (d) 1 month later
(c) (d) Photos courtesy M. Edelstein.
writings with considerable understanding. During the on the anatomy of the graft union. Development of
days of the Roman Empire, grafting was very popular, some of the early grafting techniques have been
and methods were precisely described in the writings of reviewed by Wells (178).
that era. Paul the Apostle, in his Epistle to the Romans, Liberty Hyde Bailey in The Nursery Book (8), pub-
discussed grafting between the “good” and the “wild” lished in 1891, described and illustrated the methods of
olive trees (Romans 11:17–24). grafting and budding commonly used in the United
The Renaissance period (AD 1350–1600) saw a States and Europe at that time. The methods used today
renewed interest in grafting practices. Large numbers differ very little from those described by Bailey.
of new plants from foreign countries were imported
into European gardens and maintained by grafting.
By the 16th Century, the cleft and whip grafts were
TERMINOLOGY
widely used in England and it was realized that the cam- Grafting is the art of joining grafting The union
bium layers must be matched, although the nature of two pieces of living plant of a root system
this tissue was not then understood or appreciated. tissue together in such a (understock) with
Propagators were handicapped by a lack of a good manner that they will unite a shoot system
grafting wax; mixtures of wet clay and dung were and subsequently grow and (scion) in such a
used to cover the graft unions. In the 17th Century, develop as one composite manner that they
orchards in England were planted with budded and plant. As any technique that subsequently grow
grafted trees. will accomplish this could and develop as
Early in the 18th Century, Stephen Hales, in his be considered a method of one composite
studies on the “circulation of sap” in plants, approach- grafting, it is not surprising (compound) plant.
grafted three trees and found that the center tree stayed that innumerable procedures
alive even when severed from its roots. Duhamel studied for grafting are described in the literature. Through the
wound healing and the uniting of woody grafts. The years, several distinct methods have become established
graft union at that time was considered to act as a type that enable the propagator to cope with almost any
of filter that changed the composition of the sap flow- grafting problem. These are described in Chapter 12
ing through it. Thoüin (163), in 1821, described 119 with the realization that there are many variations of
methods of grafting and discussed changes in growth each, and that there are other forms that can give simi-
habit resulting from grafting. Vöchting (171), in the lar results. Figure 11–5 illustrates a grafted plant and
late 19th Century, continued Duhamel’s earlier work the parts involved in the graft.
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Figure 11–5
In grafted plants the shoot system
consists of growth arising from one
(or more) buds on the scion. The
root system consists of an extension
of the original rootstock. The graft
union remains at the junction of
the two parts throughout the life of
the plant.
budding A form of Budding is a form The rootstock (understock, stock) is the lower
grafting that uses a of grafting. However, the portion of the graft, which develops into the root system
smaller scion piece— scion is reduced in size of the grafted plant. It may be a seedling, a rooted cut-
sometimes just a piece and usually contains only ting, a layered or micropropagated plant. If the grafting
of the stem with an one bud. An exception to is done high in a tree, as in topworking, the rootstock
axillary bud. this is patch budding of may consist of the roots, trunk, and scaffold branches.
pecan, where secondary The interstock (intermediate stock, interstem)
and tertiary buds are adjacent at the same node to the is a piece of stem inserted by means of two graft unions
primary bud. The various budding methods are between the scion and the rootstock. Interstocks are
described in Chapter 13. used to avoid incompatibility between the rootstock
The scion becomes the new shoot system of the and scion, to produce special tree forms, to control dis-
graft. It is composed of a short piece of detached shoot ease (e.g., fire-blight resistance), or to take advantage of
containing several dormant buds, which, when united their growth-controlling properties.
with the rootstock, comprises the upper portion of the Vascular cambium is a thin tissue located between
graft. The stem, or branches, or both, of the grafted the bark (periderm, cortex, and phloem) and the wood
plant will grow from the scion. The scion should be of (xylem) (see Fig. 11–6). Its cells are meristematic; that
the desired cultivar and free from disease. is, they are capable of dividing and forming new cells.
Figure 11–6
Top: Grafting terminology of the “bark”
and “wood” and associated tissues with
schematic drawing of a stem cross section
of a young woody plant stem. Bottom:
Schematic longitudinal section of the stages
of graft union formation: (Stage 1) Lining up
vascular cambiums of the rootstock and
scion, and (Stage 2) subsequent wound
healing response. (Stage 3) Callus bridge
formation. (Stage 4) Wound-repair xylem
and phloem occur in the callus bridge just
prior to initial cambium formation. (Stage 5)
The vascular cambium is completed across
the callus bridge and is forming secondary
xylem and phloem.
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For a successful graft union, it is essential that the cam- Utilization and Propagation
bium of the scion be placed in close contact with the of Clonal Rootstock
cambium of the rootstock. Clonal rootstocks are those vegetatively propagated by
callus Tissue
Callus is a term applied stool layering, rooted cuttings, or micropropagation.
composed of
to the mass of parenchyma Micropropagation of clonal rootstocks makes possible
parenchyma cells,
cells that develop from and the production of great numbers of such plants, upon
which is a response
around wounded plant tis- which the scion cultivar can be grafted or budded
to wounding. Callus
sues. It occurs at the junction (76, 77). Rootstock of citrus is produced from apomic-
development is
of a graft union, arising from tic seed and is genetically uniform; this is a more cost-
important in graft
the living cells of both the effective method of propagating clonal rootstock than
union formation.
scion and rootstock. The traditional asexual techniques.
production and interlocking Clonal rootstocks are desirable not only to pro-
of these parenchyma (or callus) cells constitute one of duce uniformity, but also to utilize special characteris-
the important steps in callus bridge formation between tics such as disease resistance. Clonal rootstock also
the scion and rootstock in a successful graft. influence the size and growth habit of the grafted plant
and flowering and fruit development of the scion. Each
particular scion-rootstock combination requires an
SEEDLING AND CLONAL
extensive evaluation period in different environments
ROOTSTOCK SYSTEMS before its future performance can be predicted.
Rootstocks can be divided into two groups: seedling Historically, clonal rootstocks for fruit crops
and clonal. received much attention in European and Middle
Eastern countries, going back centuries. Today, much
of the apple production around the world is on clonal
Utilization and Propagation
rootstocks for size control and fruit yield. Other fruit
of Seedling Rootstock
crops, such as pear, quince, plum, cherries, grapes, cit-
Seedling rootstocks propagated from seed can be mass- rus, and others are routinely propagated on clonal root-
produced relatively simply and economically. Viruses stock (179).
are transmitted from parent to progeny in very low Only pathogen-free scions and rootstock material
percentages or not at all except in specific instances. should be utilized in the nursery. To maintain rootstock
Seedling plants tend to have deeper rooted and more influence, deep planting of the nursery tree or grafted
firmly anchored plants than rootstocks grown from vegetable—which may lead to “scion rooting”—must
cuttings (e.g., plum and apple rootstock). be avoided, as illustrated in Figure 11–7. The deeper
Seedling rootstock may show genetic variation the graft union below the soil surface, the higher the
leading to variability in growth and performance of the incidence of scion rooting is likely to be (31).
scion variety. The variation can arise from natural het-
erozygosity of the source or from cross-pollination—both
are more likely if the rootstock is from an unknown, REASONS FOR GRAFTING
unselected source. Selection of special mother-tree AND BUDDING
(elite) seed source trees or a special clone can provide
Grafting and budding serve many different purposes:
uniform, special seedling rootstocks for specific crops
(see Chapter 5). • Perpetuating clones desired for their fruiting, flower-
Uniformity of seedling variability can be con- ing, or growth characteristics that cannot be readily
trolled by managing production conditions in the nurs- maintained or economically propagated by other
ery, including digging nursery trees of the same age, asexual means
one row at a time, and discarding off-type or slow- • Combining different cultivars into a composite plant
growing seedlings or budded trees. In most nurseries, as scion, rootstock, and interstock—each part pro-
the young trees are graded by size, and all those of the viding a special characteristic
same grade are sold together. Many fruit crops grown • Changing cultivars of established plants (topwork-
on uniform seedling rootstocks show no more variabil- ing), including combining more than one scion culti-
ity resulting from the rootstock than from unavoidable var on the same plant
environmental differences in the orchard—principally • Repairing graftage for injuries—including inarching
soil variability. and bridge graftage
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(b)
Figure 11–9
Grafted ornamental (a) cactus and (b) succulents. An easily rooted cultivar is used as
the rootstock and an unusual attractive type is used as the scion. These grafts are
made in large quantities in Japan and Korea, and shipped to wholesale nurseries in
(a) other countries for rooting, potting, and growing until ready for sale in retail outlets.
abiotic stress abiotic stress, size con- to become dwarfed (Fig. 11–10). Scions grafted onto
A condition caused by trol, enhanced reproduc- selected rootstock of some citrus, pear, and apple root-
environmental factors tive growth, reduction in stocks produce larger size and/or better-quality fruit
such as drought, low nursery production time, than when grafted onto other rootstock (179).
temperature, low and increased transplant-
Hastening Reproductive Maturity. Scions of many fruit
oxygen, and salinity, ing success.
crops can be established more quickly in the orchard and
which reduce growth
come into bearing more rapidly when grafted onto dwarf-
and can sometimes kill Greater Resistance to
ing rootstock (169), as opposed to being grown as
plants. Environmental Stress
seedlings or as rooted cuttings. (An exception to this is
biotic stress and Disease. For many
peach production in Mexico, where very vigorous
A condition caused by kinds of plants, root-
seedlings are selected for fruit production—seedling
living organisms such stocks are available that
plants fruit as rapidly as grafted plants.) It is also possible
as insects, pathogens, tolerate unfavorable abi-
to hasten the onset of maturity by grafting cultivars onto
and nematodes that otic stress conditions—
larger, established trees. Such grafting takes advantage of
reduce growth and can such as heavy, wet soils,
an existing large root system of the rootstock plant to
sometimes kill plants. salinity, and drought
speed up maturation of the scion.
(Figs. 11–1, 11–2, and
11–4) (47, 124–126, 129). Other rootstocks may resist Hastening Plant Growth Rate and Reducing Nursery
biotic stresses such as soil-borne insect, nematodes, Production Time. In nursery production of shade
viruses, or pathogens (34, 86) better than the plant’s own trees, budded or grafted trees grow more rapidly than
roots (Fig. 11–3). See Chapters 19 and 20 for detailed seedling or cutting-produced trees; for example, Acer
discussions of the rootstocks available for the various platanoides ‘Crimson King’ budded on a vigorous root-
fruit and ornamental species. Special rootstocks for stock (see Fig. 13–4), and budded Tilia cordata or bud-
glasshouse, poly-covered high tunnel production and ded Zelkova serrata grow more in 1 year than rooted
field production of vegetable crops are used in Europe, cuttings will in 3 or 4 years (53).
the Middle East, Asia, and North America to avoid Improving Transplanting Success. Some plants
root diseases such as Monosporascus, Fusarium and rooted by cuttings make such poor root systems that
Verticillium wilt (34, 131). In the Netherlands, green- they are difficult or impossible to transplant; for
house cucumbers are grafted onto Cucurbita ficifolia, example, the Koster spruce (Picea pungens) can be
and commercial tomato cultivars are grafted onto vig- rooted in commercial numbers, but cannot be suc-
orous F1 hybrid, disease-resistant rootstocks (21). cessfully transplanted unless the root system is pro-
Controlling Size of Grafted Plant. For some species, duced from grafted plants (53). Many Asiatic maples
size-controlling rootstocks are available that can cause form poor root systems from cuttings and must be
the composite grafted plant to have exceptional vigor or grafted (170).
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Figure 11–10
Relative size of apple trees on different rootstock. The reduction in tree size ranges from dwarfing (25 to 50 percent of a
standard full-size tree) to semi-dwarfing (60 to 70 percent) to vigorous to very vigorous (same size as a seedling tree). With the
exception of Antonovka, all listed are clonal rootstock. The absolute size of the mature, composite tree is determined by soil,
climate, culture, and the vigor of the scion cultivar (e.g., the scions of the vigorous cultivar ‘Mutsu’ are twice as large as ‘Golden
Delicious’ on ‘Malling 9’ dwarfing rootstock).
Obtaining the Benefits of Certain Interstocks • Obtaining special forms of plant growth. By grafting
(Double-Working) In addition to the rootstock and certain combinations together it is possible to pro-
scion, one may insert a third plant system between duce unusual types of plant growth, such as “tree”
them by grafting. Such a section is termed an roses (Fig. 11–12) or “weeping” cherry, birch, or wil-
interstock, interstem, intermediate stock, or low cultivars (Fig. 11–8).
intermediate stem section. This is done by making Nurseries supplying trees on seedling or clonal
two grafts (see Fig. 12–50), or double budding. For rootstocks, or with a clonal interstock, should identify
example, a thin plate (minus the bud) of ‘Old Home’ such stocks on the label just as they do for the scion
pear interstock is budded on the quince rootstock, cultivar.
then a shield bud of the ‘Bartlett’ scion is inserted
directly over the ‘Old Home’ plate and wrapped with
a budding rubber (see Changing Cultivars of Established
double-working The Plants (Topworking)
Fig. 13–21).
grafting or budding of
There are several A fruit tree, or an entire orchard, may be replaced with a
an interstock (interstem)
reasons for using double- more desirable cultivar. It could be unproductive, or an
between the rootstock
working in propagation: old cultivar whose fruits are no longer in demand; it
and scion.
could be one with poor growth habits, or possibly one
• The interstock makes it possible to avoid certain that is susceptible to topworking
kinds of incompatibility. prevalent diseases or The grafting of a new
• The interstock may possess a particular characteristic insects. Topworking cultivar onto established
(such as disease resistance or cold-hardiness) not pos- has sometimes been trees in the orchard.
sessed by either the rootstock or the scion. done by California pro-
• A certain scion cultivar may be required for disease ducers of peach, plum, and nectarine every 2 to 3 years
resistance in cases where the interstock characteristics to take advantage of newer, more promising cultivars
are the chief consideration, such as in the control of and thus remain competitive on the market. Examples
leaf blight on rubber trees (Hevea) (84). of topworking are shown in Figure 11–13, page 424.
• The interstock may reduce vegetative growth and In an orchard of a single cultivar of a species requir-
enhance reproductive growth of the tree. For exam- ing cross-pollination, provision for adequate cross-
ple, when a stem piece of the dwarfing ‘Malling 9’ pollination can be obtained by topworking scattered trees
apple rootstock is used as an interstock and inserted throughout the orchard to a proper pollinating cultivar.
between a vigorous rootstock and a vigorous scion A single pistillate (female) plant of a dioecious (pistillate
cultivar, it reduces growth of the composite tree and and staminate flowers borne on separate individual
stimulates flowering and fruiting in comparison plants) species, such as the hollies (Ilex), may be unfruit-
with a similar tree propagated without the interstock ful because of the lack of a nearby staminate (male) plant
[Fig. 11–11 (132).] to provide proper pollination. This problem can be
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(a) (b)
Figure 11–11
Effect of interstock on the size
of six-year-old ‘Cox’s Orange
Pippin’ apple scion grafted
on a vigorous ‘MM 104’
rootstock: (a) Cox/’M 9’
dwarfing interstock/’MM 104’,
(b) Cox/’M 27’ dwarfing
interstock/’MM 104’,
(c) Cox/’MM 104’ vigorous
interstock/’MM 104’,
(d) Cox/’M 20’ dwarfing
(c) (d) interstock/’MM 104’.
(b)
Figure 11–13
TopWorking. (a) Inlay bark graft in
top working an orchard. (b) Top
worked citrus grove in Sicily using an
inlay bark graft. (c) Smaller citrus
(a) (c) liner with inlay bark graft.
grafting, or inarching, such damage can be repaired and another technique used to clean up viruses and bacterial
the tree saved. This is discussed in detail in Chapter 12. problems with budwood (112).
stands of forest species of pine, hemlock, oak, and graft interface or callus bridge, and vascular differen-
Douglas-fir (59, 97). Such root grafts are common tiation across the graft interface (106).
between roots of the same tree or between roots of The scion will not resume its growth successfully
trees of the same species. Grafts between roots of trees unless a vascular connection has been established so
of different species are rare. In the forest, living that it may obtain water and mineral nutrients.
stumps sometimes occur, kept alive because their Likewise, degeneration of the rootstock will occur if the
roots have become grafted to those of nearby intact, phloem in the graft union is disrupted from sending
living trees, allowing the exchange of nutrients, water, carbohydrates and other metabolites from the scion to
and metabolites (95, 97). the root system. In addition, the scion must have a ter-
The anatomy of natural grafting of aerial roots minal meristematic region—a bud—to resume shoot
has been studied (128). Natural root grafting also per- growth and, eventually, to supply photosynthate to the
mits transmission of fungi, viruses, and phytoplasmas root system.
from infected trees to their neighbors (128). This prob- Considering in more detail the steps involved in
lem can occur in orchard and nursery plantings of trees graft union formation (Figs. 11–6 and 11–14), the
and in urban shade tree sites where numerous root first one listed below is a preliminary step, but never-
grafts may result in the slow spread of pathogens theless, it is essential, and one over which the propaga-
throughout the planting. Natural root grafting is a tor has control.
potential source of error in virus-indexing procedures
where virus-free and virus-infected trees are grown in
close proximity (60). In addition, fungal pathogens
causing oak wilt and Dutch elm disease can be spread
by such natural root connections.
FORMATION OF THE
GRAFT UNION
A number of detailed studies have been made of graft
union formation, with woody (9, 11, 35, 49, 133, 156,
168) and herbaceous plants (52, 91, 101, 105, 123,
152, 159, 164, 188). Just as de novo meristems are
necessary for adventi-
de novo meristems
tious bud and root
New meristematic
formation, a de novo-
areas initiated from
formed meristematic
parenchyma cells such
area (new vascular cam-
as the vascular cambium
bium) must develop
that must develop in
between the scion and
the callus bridge of a
rootstock if successful
grafted plant.
graft union formation is
to occur (188). The parts of the graft that are originally
prepared and placed in close contact do not themselves
move about or grow together. Rather, the union is
accomplished entirely by cells that develop after the
actual grafting operation has been made. The graft
union is initially formed by rapidly dividing callus cells,
originating from the scion and rootstock, which later
differentiate to form the vascular cambium (a lateral Figure 11–14
Graft union formation in grafted pea roots (91, 159). This
meristem) and the associated vascular system.
sequence of grafting events is common to topgrafting and
The development of a compatible graft is typi- root grafting in many other woody and herbaceous plant
cally comprised of three major events: adhesion of the species. What will vary is the time period in grafting events
rootstock and scion, proliferation of callus cells at the with different species.
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(a)
Figure 11–18
Figure 11–17 Accumulation of dictyosomes along the cell walls adjacent to
Cross section of a Hibiscus wedge graft showing the the necrotic layer at six hours after grafting in the compatible
importance of callus development in the healing of a graft autograft in Sedum telephoides. ×17,500. Courtesy R. Moore and
union. Cambial activity in the callus has resulted in the D. B. Walker (101).
production of secondary tissues that have joined the vascular
tissues of the stock and scion ×10. Photo courtesy K. Esau.
sc
rs
(1)
w
w
p
w
(2) (3) ER
Figure 11–19
Schematic diagram of secondary (de novo) formation
GV of plasmodesmata at the graft interface (callus bridge).
(1) Approaching callus cells of scion (sc) and rootstock
(rs). Pectic material (p) between adjoining callus cell
walls. Region between arrows: wall parts where
secondary plasmodesmata will be formed, as shown in
(4) (5) detail. Formation of continuous cell connections (2 to 7)
GV ER by plasmalemma and endoplasmic reticulum (ER) fusion
of adjoining cells (5, 6) within wall parts that have been
w thinned synchronously with both cell partners.
Elongation of the branched and single strands during
w rebuilding the modified wall parts (6, 7). W = cell wall,
GV = golgi vesicles, *new deposited wall material.
(6) (7) ER Redrawn from Kollman and Glockmann (80).
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The wound-repair wound-repair xylem wound cambium differentiates prior to the bridg-
xylem (wound-type (Wound-type vascular ing of vascular tissue, and in autografts of Sedum
vascular elements) elements) generally the (Crassulaceae) where procambial differentiation
is generally the first first differentiated tissue occurs before vascular differentiation (101). With
differentiated tissue to bridge the graft budding, the scion is considerably smaller and nor-
to bridge the graft union, followed by mally limited to one bud and a short shoot piece;
union, followed by wound-repair phloem. hence, any early vascular differentiation from callus
wound-repair cells is probably limited by lower phytohormone
phloem (Fig. 11–20). Initial xylem tracheary ele- levels. The vascular cambium can form independ-
ments and, frequently, initial phloem sieve tubes ent of any xylem or phloem (28), or the cambium
form directly by differentiation of callus into these may differentiate between the wound-bridging
vascular elements. A vascular cambium layer subse- xylem and phloem (159). It is important that the
quently forms between the vascular systems of the vascular cambium unite so that the continuity of
scion and rootstock. wound-bridging xylem and phloem can be main-
Exceptions to this developmental sequence are tained, and so that secondary vascular development
in bud graftage in citrus, apple, and rose where a occurs for successful graft union formation.
(a)
(b)
Figure 11–20
Vascular connections between melon and Cucurbita rootstock. (a) Early
vascular strands in callus bridge area which are from wound-repair xylem
and wound-repair phloem. (b) Vascular connections after 14 days.
(c) Schematic of vascular connections (dotted red lines between scion
(c) and rootstock. Courtesy M. Edelstein.
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Figure 11–21
Schematic of tip grafting of cactus. Top: In Method I,
the vascular bundles of the scion and rootstock were
placed together, or 1 mm (Method II) or 3 mm
(Method III) apart. Bottom: Auxin in lanolin paste
promoted vascular connections between misaligned
graft partners and increased the diameter of the
connecting vascular bundle. Redrawn from Shimomura and
Fuzihara (152).
At the edges of the newly formed callus mass, bridge, until a continuous cambial connection
parenchyma cells touching the cambial cells of the forms between rootstock and scion.
rootstock and scion differentiate into new cam- 5. Production of Secondary Xylem and Phloem
bium cells within 2 to 3 weeks after grafting. This from the New Vascular Cambium in the Callus
cambial formation in the callus mass proceeds far- Bridge
ther and farther inward from the original rootstock The newly formed cambial layer in the callus
and scion cambium, and on through the callus bridge begins typical cambial activity, laying down
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 432
new secondary xylem toward the inside and cambial zone remains attached to the inside of the
phloem toward the outside. bark flaps. Very shortly after the bud shield is
In the formation of new vascular tissues fol- inserted, a necrotic plate or layer of material develops
lowing cambial continuity, the type of cells formed from the cut cells. Next, after about two days, callus
by the cambium is influenced by the cells of the parenchyma cells start developing from the rootstock
graft partners adjacent to the cambium. For exam- xylem rays and break through the necrotic plate.
ple, xylem ray cells are formed where the cambium Some callus parenchyma from the bud scion ruptures
is in contact with xylem rays of the rootstock, and through the necrotic area in a similar manner. As
xylem elements where they are in contact with additional callus is produced, it surrounds the bud
xylem elements (122). shield and holds it in place. The callus originates
Production of new xylem and phloem thus almost entirely from the rootstock tissue, mainly
permits the vascular connection between the scion from the exposed surface of the xylem cylinder. Very
and the rootstock. It is essential that this stage be little callus is produced from the sides of the bud
completed before much new leaf development shield (scion).
arises from buds on the scion. Otherwise, the Cell proliferation continues rapidly for 2 to
enlarging leaf surfaces on the scion shoots will have 3 weeks until all internal air pockets are filled with
little or no water to offset that which is lost by callus. Following this, a continuous cambium is estab-
transpiration, and the scion quickly will become lished between the bud and the rootstock. The callus
desiccated and die. It is possible, however, even then begins to lignify, and isolated xylem tracheary
though vascular connections fail to occur, that elements appear. Lignification of the callus is com-
enough translocation can take place through the pleted between 5 to 12 weeks after budding (108,
parenchyma cells of the callus to permit survival of 172). The developmental stages and time intervals for
the scion. In grafts of vanilla orchid, a monocot, graft union formation in T-budded citrus are listed in
scions survived and grew for 2 years with only Box 11.4.
union of parenchyma cells; however, the grafted
plants did not survive when subjected to transpira- More Rapid Union Development
tional stress (111). in Chip Budding
Anatomical studies (155) have been made comparing
graft union formation in T- and chip budding. Early
GRAFT UNION FORMATION union formation between ‘Lord Lambourne’ apple
scion and ‘Malling 26’ dwarfing rootstock showed a
IN T- AND CHIP BUDDING more rapid and complete union of xylem and cambial
bark (In grafting) In T-budding, the bud tissues of the scion and rootstock after chip budding
composed of tissues piece usually consists of compared to T-budding. This is probably due to a
from the periderm, the “bark” (periderm, much closer matching of the scion tissue to the root-
cortex, phloem, and cortex, phloem, cambium), stock stem (Fig. 11–22). Also in T-budding, the cam-
vascular cambium. and often some “wood” bium of the rootstock is lifted in the flap of “bark,” so
wood (In grafting) (xylem tissue). Attached considerable callus in-filling and development of new
composed of externally to this is a cambium must occur. There is more flexibility in chip
secondary xylem with lateral bud subtended, budding, which can be done over longer periods on
some pith (in younger perhaps, by a leaf petiole. either an active or dormant rootstock, than T-budding,
woody plants). In budding, this piece of which requires an active rootstock. In part this advan-
tissue is laid against the tage to chip budding is due to less callus filling being
exposed xylem and cam- needed, and because there is no requirement for an
bium of the rootstock, as shown diagrammatically in active cambium to lift the flap of rootstock bark, as
Figure 11–22. there is with T-budding.
Detailed studies of the grafting process in T-budding The previously mentioned advantages of chip
have been made for the rose (28), citrus (93, 94), and budding compared with T-budding have also been
apple (108). demonstrated with ‘Crimson King’ maple on Acer pla-
In the apple, when the flaps of bark on either tanoides rootstock, ‘Conference’ pear on ‘Quince A’
side of the “T” incision on the rootstock are raised, rootstock, and ‘Rubra’ linden on Tilia platyphyllos
separation occurs from the young xylem. The entire rootstock.
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Figure 11–22
(a) Tissues involved in healing of an
inserted T-bud as prepared with the
“wood” (xylem) attached to the scion
bud piece. Graft union formation occurs
when callus cells developing from
the young xylem of the rootstock
intermingle with callus cells forming
from exposed cambium and young
xylem of the T-bud piece. As the bark is
lifted on the rootstock for insertion of
(a)
the bud piece it detaches by separation
of the youngest xylem and cambial cells.
(b) A cross section of a chip bud (CB),
T-bud (TB), and rootstock (RS). Because
the chip bud substitutes exactly for the
part of the rootstock that is removed,
the cambium of the roots and scion are
placed close together, resulting in a
rapid and strong union. When a T-bud
(right) is slipped under the “bark,” the
cambium of the rootstock and scion are
not adjacent, and the initial union
formation can be weak and slow. Redrawn
(b) from B. H. Howard (68).
FACTORS INFLUENCING GRAFT influence the healing of graft unions. Factors that influ-
ence graft union success include:
UNION SUCCESS
As anyone experienced in grafting or budding knows, • Incompatibility
the results are often inconsistent. An excellent percent- • Plant species and type of graft
age of “takes” occur in some operations, but in others • Environmental conditions during and following
the results are disappointing. A number of factors can grafting
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 434
Incompatibility
One of the symptoms of incompatibility in grafts
between distantly related plants is a complete lack, or a Figure 11–23
very low percentage, of successful unions. Incompatibility Some species form profuse callusing
is discussed in greater detail starting on page 441. Grafts (arrow), which helps increase graft union
between some plants known to be incompatible, initially success. Pear is easily grafted by a whip-
will make a satisfactory union, even though the combina- and-tongue graft.
tion eventually fails.
grafting certain stone fruits, such as peaches and apri-
Plant Species and Type of Graft cots, requires more care and attention to detail.
Some plants—including hickories, oaks, and beeches— Strangely enough, grafting peaches to some other com-
are much more difficult to graft than others even when patible species, such as plums or almonds, is more suc-
no incompatibility is involved. Nevertheless, such cessful than reworking them back to peaches. One
plants, once successfully grafted, grow very well with a method of grafting may give better results than another,
perfect graft union. In grafting apples, grapes, and pears or budding may be more successful than grafting, or
(Figs. 11–23 and 11–24), even the simplest techniques vice versa. For example, gymnosperms are grafted,
usually give a good percentage of successful unions, but whereas many angiosperm cultivars tend to be budded,
(a)
(c) (d)
Figure 11–24
A high take occurs when grapes are saddle grafted, but the same graft is unsuccessful with roses, which did
not form sufficient callus. (a) Heitz saddle graft bench graft tool. (b) Unsuccessful saddle graft with rose.
(b) (c and d) Successful saddle-grafted grape with profuse callusing in the callus bridge area.
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 435
with no provision for resisting desiccation. If exposed Initiation of cambial activity in the spring results
to drying air they will be killed. This was found to be from the onset of bud activity, because shortly after the
the case in studies of the effect of humidity on the heal- buds start growth, cambial activity can be detected
ing of apple grafts. Air moisture levels below the satura- beneath each developing bud, with a wave of cambial
tion point inhibited callus formation; desiccation of activity progressing down the stems and trunk. This
cells increased as the humidity dropped. In vitro studies stimulus is due, in part, to production of auxin originat-
(43) of stem pieces of ash (Fraxinus excelsior) have ing in the expanding buds (175). Callus proliferation—
shown that callus production on the cut surfaces was essential for a successful graft union—occurs most read-
markedly reduced as the water potential decreased. ily at the time of year just before and during “bud-break”
Water is one of the driving forces for cell enlarge- in the spring, because auxin gradients diminish through
ment and is necessary for callus bridge formation the summer and into fall. Increasing callus proliferation
between the stock and scion. Water must be utilized takes place again in late winter, but this is not dependent
initially from scion tissue, and if below a certain water upon the breaking of bud dormancy.
potential, insufficient water is available for callus for- When T-budding seedlings in the nursery in late
mation. Failed grafts of well-hydrated Sitka spruce summer, it is important that they have an ample supply
rootstocks produced no callus at the graft union, sug- of soil moisture just before and during the budding oper-
gesting that callus formation at the cut surface is con- ation. If they should lack water during this period, active
trolled or dependent on the formation of callus from growth is checked, cell division in the cambium stops,
the scion (10). Until vascular connections are formed and it becomes difficult to lift the bark flaps to insert the
between the rootstock and scion, the callus bridge pro- bud. At certain periods of high growth activity in the
vides the initial pathway for water, bypassing damaged spring, plants exhibiting strong root pressure (such as
xylem vessels and tracheids of the scion and rootstock. the walnut, maple, and grape) show excessive sap flow or
Within the first 3 to 4 days of callus bridge formation, “bleeding” when cuts bleeding A process in
there is a recovery of scion water potential (10); with are made preparatory to which a plant has strong
maturation of the connecting tracheids, water potential budding and grafting. root pressure that
and osmotic potential continue to increase (15, 16). Grafts made with such causes excess sap flow
Photosynthesis declines and does not increase until moisture exudation that can reduce grafting
xylem connections become reestablished (18). around the union will success.
Unless the adjoining cut tissues of a completed not heal properly. Such
graft union are kept at a very high humidity level, the “bleeding” at the graft union can be overcome by mak-
chances of successful healing are poor. With most ing slanting knife cuts below the graft around the tree.
plants, thorough waxing of the graft union or sealing of Cuts should be made through the bark and into the
the graft union with polyethylene grafting tape, xylem to permit such exudation to take place below the
Parafilm, or Buddy Tape (Aglis & Co. Ltd.) helps retain graft union. Containerized rootstock plants of Fagus,
the natural moisture of the tissues, which is all that is Betula, or Acer are relocated to a cool place with reduced
necessary. Often root grafts are not waxed but stored in watering until the “bleeding” stops. Then plants are
a moist (not overly wet) packing material during the grafted after the excessive root pressure subsides.
callusing period. Slightly damp peat moss or wood On the other hand, dormant containerized root-
shavings are good media for callusing, providing ade- stocks of junipers or rhododendrons, when first brought
quate moisture and aeration. into a warm greenhouse in winter for grafting, should
be held for several weeks at 15 to 18°C (60 to 65°F)
Growth Activity of the Rootstock until new roots begin to form. Then the rootstocks are
Some propagation methods, such as T-budding and bark physiologically active enough to be successfully grafted.
grafting, depend on the bark “slipping,” which means When the rootstock is physiologically overactive
that the vascular cambium is actively dividing, producing (excessive root pressure and “bleeding”), or underactive
young thin-walled cells on each side of the cambium. (no root growth),
These newly formed cells separate easily from one some form of side graft top-grafting A form of
another, so the bark “slips” (Fig. 11–22). Chip budding can be used, in which grafting in which the shoot
can be done on a dormant or active rootstock. Hence, the rootstock top is of the rootstock is com-
there is much more flexibility in scheduling chip bud- initially retained. On pletely removed at the time
ding, because there is no requirement for an active cam- the other hand, top- the graft is made (e.g.,
bium to lift the flap of rootstock bark, as with T-budding. grafting, in which in-lay bark graft of pecan).
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 437
Figure 11–26
Polarity in grafting. In topgrafting, the proximal end of the scion is
attached to the distal end of the rootstock. In root grafting,
however, the proximal end of the scion is joined to the proximal
end of the rootstock.
the top of the rootstock is completely removed at the In nurse-root grafting, the graft union is purposely set well
time the graft is made, is likely to be successful in plants below the ground level, and the scion itself produces
in which the rootstock is neither overactive nor underac- adventitious roots, which ultimately become the entire
tive (44). root system of the plant. See Figure 12–26, page 487 for
greater detail of nurse-root grafting systems.
Polarity in Grafting In T-budding or patch budding, the rule for
observance of correct polarity is not as exacting. The
Distal and Proximal Ends Correct polarity is strictly
buds (scion) can be inserted with reversed polarity and
observed in commercial grafting operations. As a gen-
eral rule, (and as shown in Fig. 11–26), in top-grafting,
the proximal end of the scion should be inserted into
the distal end of the rootstock. But in normal root graft-
ing, the proximal end of the scion should be inserted
into the proximal end of the rootstock.
Should a scion be inserted with reversed polar-
ity “upside-down,” it is possible for the two graft
unions to be successful and the scion to stay alive for
a time (Fig. 11–27). But in bridge grafting, the
reversed scion does not increase from its original size,
whereas the scion with correct polarity enlarges nor-
mally (Fig. 11–28).
Nurse-Root Grafting Nurse-root grafting is a
temporary graft system to allow a difficult-to-root plant
to form its own adventitious roots. The rootstock may be
turned upside-down, its polarity reversed, and then
grafted to the desired scion. A temporary union will form,
Figure 11–27
and the rootstock will supply water and mineral nutrients Inverse graft of grape with graft union forming between the
to the scion, but the scion is unable to supply necessary distal end of the scion to the distal end of the rootstock.
organic materials to the rootstock, which eventually dies. Notice that the shoot reorients itself via gravitational response.
the resulting plant (121). In stone fruit propagation, halfway through the rootstock shoot above the bud union
bud-wood, free of ring-spot virus, has consistently and breaking the shoot over the rootstock stem), girdling,
greater “takes” than infected bud-wood. or totally removing the rootstock above the scion bud
Top-grafting olives in California is seriously hin- union, apical dominance is broken and the scion bud rap-
dered in some years by attacks of the American plum idly elongates (Fig. 11–30) (50).
borer (Euzophera semifuneralis), which feeds on the soft With budded citrus, plants on which rootstock
callus tissue around the graft union, resulting in the shoots remained attached (lopping, or bending the
death of the scion. In England, nurseries are often rootstock shoot to its base and tying it in position)
plagued with the red bud borer (Thomasiniana had the greatest gains in scion growth. This was due
oculiperda), which feeds on the callus beneath the bud- to the greater transfer of photosynthate from the root-
shield in newly inserted T-buds, causing them to die. stock leaves to scion shoots during growth flushes,
and to roots during periods between growth flushes
Plant Growth Regulators and Graft (181, 182).
Union Formation
Plant growth regulators, particularly auxin, applied to
tree wounds or to graft unions give variable results in GENETIC LIMITS OF GRAFTING
wounding response and graft union formation (93, 118,
Since one of the requirements for a successful graft
152). Auxin (IBA, NAA) and cytokinin (BA) enhance
union is the close matching of the callus-producing
graft success when applied to the base of side-grafted
tissues near the cambium layers, grafting is generally
Picea scions, while the plant growth retardant, dikegulac,
confined to the dicotyledons in the angiosperms, and
stimulated scion growth by retarding rootstock develop-
to gymnosperms. Both have a vascular cambium layer
ment (17). Cytokinins enhance patch budding of
existing as a continuous tissue between the xylem and
Persian walnut. The eloquent work of Shimomura (152)
the phloem. Grafting is more difficult, with a low per-
in tip grafting of cactus demonstrated how auxins
centage of “takes” in monocotyledonous plants.
enhanced vascular connections of deliberately mis-
Monocots have vascular bundles scattered throughout
aligned scions (Fig. 11–21). TIBA, a well-known
the stem, rather than the continuous vascular cam-
inhibitor of basipetal transport of auxin, inhibited vascu-
bium of dicots. However, there are cases of successful
lar connections in the graft union; however, by subse-
graft unions between monocots. By making use of the
quent reapplication of auxin, the inhibitory effect of
meristematic properties found in the intercalary tis-
TIBA was eliminated and vascular connections occurred.
sues (located at the base of internodes), successful
However, unlike auxin usage in cutting propaga-
grafts have been obtained with various grass species as
tion, no plant growth regulators are routinely used in
well as the large tropical monocotyledonous vanilla
commercial grafting and budding systems. In general,
orchid (111).
plant growth regulators do not uniformly enhance
Before a grafting operation is started, it should be
grafting, nor do they overcome graft incompatibility.
determined that the plants to be combined are capable
of uniting and producing a permanently successful
Post-Graftage—Bud-Forcing Methods union. There is no definite rule that can exactly predict
After graft union formation has occurred in grafting or the ultimate outcome of a particular graft combination
budding, it is often necessary to force out the scion or the except that the more closely the plants are related
scion bud. In field budding of roses, 2 to 3 axillary buds botanically, the better the chances are for the graft
of the rootstock remain distal to the scion bud. The union to be successful (71). However, there are numer-
axillary buds of the rootstock, which develop into photo- ous exceptions to this rule.
synthesizing branches,
crippling The bending are initially important
(restriction) or cutting for the growth of the Grafting Within a Clone
halfway through the composite plant. But A scion can be grafted back onto the plant from which
rootstock stem above they can inhibit growth it came, and a scion from a plant of a given clone can
the bud union to helps of the scion through be grafted onto any other plant of the same clone. For
force out the bud and apical dominance, which example, a scion taken from an ‘Elberta’ peach tree
maintain growth of the is an auxin response. could be grafted successfully to any other ‘Elberta’
grafted plant. By “crippling” (cutting peach tree in the world.
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 440
(a)
(c) (d)
(b)
Figure 11–30
Forcing or “crippling” of (a and b) T-budded apples; (c and d) Chip budded roses. The rootstock is partially severed on the same
side (arrows) that the rootstock was budded. This breaks the apical dominance of the rootstock shoot system on the scion, and
helps force out the scion bud. By not totally severing the rootstock top, growth of the composite plant is maximized, since the
shock of total severance to the composite plant is avoided, and photosynthate is still produced by the rootstock (182). The
rootstock shoot system will be totally severed later, and the scion will fully develop into the shoot system of the composite plant.
Grafting Between Clones persica). But on the other hand, almond and apricot,
Within a Species both in the same genus, cannot be intergrafted success-
In tree fruit and nut crops, different clones within a fully. The ‘Beauty’ cultivar of Japanese plum (Prunus
species can almost always be grafted without difficulty salicina) makes a good union when grafted on almond,
and produce satisfactory trees. However, in some but another cultivar of P. salicina, ‘Santa Rosa,’ cannot
conifer species, notably Douglas-fir (Pseudotsuga men- be successfully grafted on almond. Thus, compatibility
ziesii), incompatibility problems have arisen in grafting between species in the same genus depends on the par-
together individuals of the same species, such as ticular genotype combination of rootstock and scion.
selected P. menziesii clones onto P. menziesii seedling Reciprocal interspecies grafts are not always
rootstock (36). Incompatibility is also a problem in successful. For instance, ‘Marianna’ plum (Prunus
grafting clones of deciduous species, such as red maple cerasifera × P. munsoniana) on peach (Prunus persica)
(Acer rubra), Chinese chestnut (Castanea mollissima), roots makes an excellent graft combination, but the
and red oak (Quercus rubra). reverse—grafts of the peach on ‘Marianna’ plum roots—
either soon die or fail to develop normally (2, 90).
Grafting Between Species
Within a Genus Grafting Between Genera
For plants in different species but in the same genus, Within a Family
grafting is successful in some cases but unsuccessful in When the plants to be grafted together are in the same
others. Grafting between most species in the genus family but in different genera, the chances of a success-
Citrus, for example, is successful and widely used com- ful union become more remote. Cases can be found in
mercially. Almond (Prunus amygdalus), apricot (Prunus which such grafts are successful and used commercially,
armeniaca), European plum (Prunus domestica), and but in most instances such combinations are failures.
Japanese plum (Prunus salicina)—all different species— Intergeneric grafts are rarely used in conifers. However,
are grafted commercially on rootstock of peach (Prunus high success rates occur between Nootka cypress
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 441
Scion
Callus
X
X
P
P X
X
(a) (b)
Figure 11–31
Graft incompatibility in ‘Jonagold’ apple scions budded to dwarfing ‘Mark’ rootstock. (a) Unstained section, with callus tissue
between the rootstock and scion. (b) Section stained with toluidine blue O. The xylem (x) in the graft union is interrupted by
parenchyma tissue (arrows) which limits water flow and survival of the scion. Courtesy of M. R. Warmund (176).
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 442
(a) (b)
Figure 11–34
Figure 11–32 Graft incompatibility occurring some 15-plus years after the
(a) Compatible apple chip bud with vascular continuity indicated Monterey pine (Pinus radiata) was grafted.
by red dye, azosulfonate. (b) Unsuccessful chip bud with vascular
discontinuity, as indicated by no visible dye. Courtesy M. R. Warmund.
in by proliferating ray tissue that does not lignify with apparent success (Figs. 11–33 and 11–34) (35)
normally (109). but gradually develop distress symptoms with time,
Incompatiblity can occur within a period of days due either to failure at the union or to the develop-
or years (Figs. 11–33 and 11–34). Delayed incompatibil- ment of abnormal growth patterns . Incompatibility of
ity can take as long as 20 years to occur with confiers and citrus and Monterey pine (Pinus radiata) may occur 15
oaks. Some apricot cultivars grafted onto myrobalan or more years after grafting (Fig. 11–34). Nelson (113)
plum rootstick will not break at the graft union until the has developed an extensive survey of incompatibility
trees are fully grown and bearing crops (46). in horticultural plants which should be consulted
The distinction between a compatible and an before attempting graft combinations between species
incompatible graft union is not always clear-cut. whose graft reactions are unknown to the grafter.
Incompatible rootstock-scion combinations can com- Other summaries of graft compatibility have been
pletely fail to unite. Frequently they unite initially published (2).
Figure 11–35
Graft compatibility affects
water uptake. (a) Arava
melon showing hotter scion
and cooler temperature in
Cucurbita rootstock (arrow)
with noncompatible grafting
combination. (b)
Compatible graft showing
uniform temperature
between scion and
rootstock. Differences in
temperature gradients
determined with a thermal
(a) (b) camera. Courtesy M. Edelstein.
• Suckering of rootstock (Fig. 11–37). roots (see Fig. 11–33), or much later with conifers and
• Graft components breaking apart cleanly at the graft oaks (Fig. 11–34). While the scion overgrowing the root-
union. stock (or rootstock outgrowing the scion) at the graft
union is not a reliable indicator, it is sometimes associated
An isolated case of one or more of the preceding
with incompatibility (Figs. 11–38 and 11–39) (2, 26).
symptoms (except for the last) does not necessarily mean
the combination is incompatible. Incompatibility is
clearly indicated by trees breaking off at the point of Anatomical Flaws Leading
union, particularly when they have been growing for to Incompatibility
some years and the break is clean and smooth, rather than With incompatible cherry (Prunus) grafts, the number
rough or jagged. This break may occur within a year or of well-differentiated phloem sieve tubes is much lower
two of the union, for instance, in the apricot on almond at and below the union. There is a greater autolysis of
cells, and generally a very low degree of phloem differ-
entiation (149). Poor differentiation of the phloem
below the union may be due to a lack of hormones, car-
bohydrates, and other factors—the size of the sieve
tubes depends on auxin, cytokinin, and sucrose levels
(149). With incompatible apricot/plum (Prunus)
grafts, some callus differentiation into cambium and
vascular tissue does occur; however, a large portion of
the callus never differentiates (Fig. 11–40) (48). The
union that occurs is mechanically weak.
With incompatible apple grafts, vascular disconti-
nuity occurs with xylem interrupted by parenchyma tissue
(Figs. 11–31 and 11–32) (176), which disrupts normal
xylem function leading to death of the budded scion.
Nontranslocatable (Localized)
Incompatibility
For lack of better terminology, physiological factors of
graft incompatibility has been traditionally classified as
Figure 11–36 nontranslocatable (localized) or translocatable
Physiological incompatibility between scion and rootstock. (109). It is difficult to distinguish differences between
Scion overgrowth caused by blockage of assimilates
the symptoms of nontranslocatable and translocatable
translocating from the scion to the rootstock, causing a weak
root system. The melon scion grafted on Cucurbita rootstock incompatibility. Anatomical symptoms of incompati-
later died as a result of insufficient support from the bility can include phloem degeneration or phloem
rootstock. Photo courtesy M. Edelstein. compression, and cambial or vascular discontinuity in
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 444
Figure 11–37
Undesirable suckering of
rootstocks. (a) Hamamelis
vernalis ‘Sandra’ grafted on
Hamamelis vernalis rootstock,
and (b) rootstock suckers on
recently grafted Ulmus alata
‘Lace Parasol’ grafted onto
seedling Ulmus alata. The
suckers will need to be
removed. Photo courtesy
(a) (b) B. Upchurch.
Figure 11–38
While rootstock outgrowth
is not desirable, a large,
strong tree can still develop.
(a) Sweet orange rootstock
used for dwarfing, overgrow-
ing the grapefruit scion.
(b) Rootstock overgrowing
scion on Morus alba
‘Platanifolia.’ Photo b courtesy
(a) (b) B. Upchurch.
Figure 11–39
Scion or rootstock outgrowth
can still lead to a large, strong
tree. Such outgrowth (arrows)
is more related to the genetic
tendency for growth, than to
incompatibility. (a) Scion
overgrowing rootstock:
Acer pentaphyllum on A.
pseudoplatanus rootstock,
and (b) grapefruit scion on
sour orange rootstock, which
tolerates alkaline, heavy soils,
but can be susceptible to
Trestiza. Photo a courtesy
(a) (b) B. Upchurch.
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:32 PM Page 445
the union area, causing mechanical weakness and sub- example of nontranslocatable incompatibility is
sequent breakdown of the union. Nontranslocatable ‘Bartlett’ (‘Williams’) pear grafted directly onto dwarf-
incompatibility includes graft combinations in which a ing quince rootstock. When mutually compatible ‘Old
mutually compatible interstock overcomes the Home’ or (‘Beurré Hardy’) is used as an interstock, the
incompatibility of the scion and rootstock. The inter- three-graft combination is completely compatible, and
stock prevents physical contact of the rootstock and satisfactory tree growth takes place (107, 122, 132).
scion and affects the physiology of the normally incom-
patible scion and rootstock. In some innovative Translocatable Incompatibility
research, membrane filters placed between graft part- Translocatable incompatibility includes certain
ners demonstrated that physical contact is not neces- graft/rootstock combinations in which the insertion of a
sary to develop compatible grafts (104, 106). A good mutually compatible interstock does not overcome
respectively, between different cells (103, 105, 106). reaction cannot be considered a universal cause of
Although incompatibility is clearly related to genetic graft incompatibility.
differences between rootstock and scion, the mecha- Phenolic compounds have also been implicated in
nisms by which incompatibility is expressed are not graft incompatibility (49). Phenolic compounds are
clear. The large number of different genotypes that can widespread in plants and present in the biochemical
be combined by grafting produces a wide range of dif- responses to stress and wounding. They play a role in lig-
ferent physiological, biochemical, and anatomical nification (27), which occurs in graft union formation.
interactions when grafted. Several hypotheses have
Modification of Cells and Tissue The lignification
been advanced in attempts to explain incompatibility.
processes of cell walls are important in the formation of
One proposed physiological and biochemical
strong unions in pear-quince grafts. Inhibition of
mechanism concerns incompatible combinations of
lignin formation and the establishment of a mutual
certain pear cultivars on quince rootstock (61). The
middle lamella results in weak graft unions. In compat-
incompatibility is caused by a cyanogenic glucoside,
ible pear-quince graft combinations, the lignin in cell
prunasin, normally found in quince but not in pear
walls at the graft union is comparable to adjacent cells
tissues. Prunasin is translocated from the quince into
outside the union (27). Conversely, adjoining cell walls
the phloem of the pear. The pear tissues break down
in the graft union of incompatible combinations con-
the prunasin in the region of the graft union, with
tain no lignin, and are interlocked only by cellulose
hydrocyanic acid (cyanide) as one of the decomposi-
fibers.
tion products (Fig. 11–42). The presence of the
With incompatible apricot-plum (Prunus) grafts,
hydrocyanic acid leads to a lack of cambial activity at
some callus differentiation into cambium and vascular
the graft union, with pronounced anatomical distur-
tissue does occur; however, a large portion of the callus
bances in the phloem and xylem at the resulting
never differentiates (Fig. 11–40) (48). The union that
union. The phloem tissues are gradually destroyed at
occurs is mechanically weak.
and above the graft union. Conduction of water and
materials is seriously reduced in both xylem and Cell Recognition of the Grafting Partners It has been
phloem. The presence of cyanogenic glycosides in woody postulated that the critical event deciding compatible and
plants is restricted to a relatively few genera. Hence, this incompatible grafts may occur when the callus cells first
Figure 11–42
Nontranslocatable incompatibility of Bartlett pear scion overcome with `old Home’ interstock on quince rootstock (61).
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 448
cellular recognition touch (189). There response to wounding. Callus proliferation is not related
The union of specific may be cellular recog- to graft compatibility-incompatibility systems, since it
cellular groups on the nition that must occur does not require a recognition event to occur; that is, cal-
surfaces of the interacting in successful graft lus proliferation occurs in wounded cuttings, as well as in
cells that results in a union formation. incompatible and compatible graft systems (101, 103).
specific defined response Alternatively, the failure Vascular differentiation in the callus bridge, which
[e.g., pollen-stigma of procambial differen- typically occurs from the severed vascular strands of the
compatibility- tiation in incompatible scion and rootstock, can occur even when the scion and
incompatibility grafts may be the result rootstock are physically separated by a porous membrane
recognition responses of a direct form of cel- filter (inserted in order to prevent direct cellular contact
with glycoprotein surface lular communication without impeding the flow of diffusible substances
receptors in flowering between the graft between the graft partners) (102, 104); this was done
plants (30)]. partners (101). with autografts of autograft The scion and
In a compatible Sedum (a herbaceous rootstock are from the
graft, the wound response is followed by a dissolution species), which may same plant or species.
of the necrotic layer, perhaps as a prerequisite to the not be representative
formation of secondary plasmodesmata between cells of graftage in woody perennial plants. Nonetheless, it is
of the graft partners (165). There is direct cellular con- evidence that successful graft union formation can occur
tact of plasmodesmata (minute cytoplamic threads that in the absence of direct cellular contact, and does not
extend through openings in cell walls and connect the require a positive recognition system.
protoplasts of connecting cells) in the callus bridge that Tissue alignment [e.g., vascular cambium of woody
symplastically connects the grafting partners plants, vascular bundles of cacti (152)] determines what
(Fig. 11–19) (81). This forms a potential communica- cell types and tissue will be differentiated in the callus
tion pathway among cells in the graft bridge, which bridge. It has been proposed that phytohormones are
may be important in cell recognition and compatibil- released from wounded vascular bundles into the
ity/incompatibility responses. surrounding tissue where they function as morphogenic
Conversely, cellular recognition may not be a fac- substances inducing and controlling the regeneration of
tor in grafting compatibility/incompatibility. Partners of cambium and vascular tissue (3). This hypothesis can be
compatible and incompatible grafts adhere during the applied to graft union formation, with phytohormones
early stages of graft union formation; this passive event such as auxin as potential morphogens needed for graft
does not require mutual cell recognition [grafted Sedum union formation. Auxin should not be considered as a
will even adhere to inert wooden objects (101, 103)], specific recognition molecule per se because of its com-
nor is it related to compatibility (106). Adhesion of graft mon occurrence and involvement in numerous other
partners results from the deposition and subsequent developmental processes (104, 106). Phytohormones
polymerization of cell wall materials that occur in (and carbohydrates, etc.), predominantly released from
the scion, enable vascular connections to develop and join heterografts, compared heterograft The scion
as a functional unit in the graft union, without any cellu- with compatible auto- and rootstock are from
lar recognition required. grafts, and adjacent root- a different cultivar or
A model for graft compatibility-incompatibility stock and scion cells species.
is presented that suggests grafts will be incompatible must produce similar
only if naturally occurring morphogens that promote lignins and have identical peroxidase enzyme patterns to
the formation of a successful graft (e.g., auxin) are ensure the development of a functional vascular system
overridden by toxins [e.g., hydrocyanic acid, ben- across the graft union (40). With electrophoresis, if the
zaldehyde (62, 63)] that elicit graft incompatibility peroxidase bands match, the combination may be com-
(Fig. 11–43) (106). patible; if they do not, incompatibility may be predicted.
There is probably no universal cause of graft Using electrophoresis is an important step in developing
incompatibility in plants (145). Most likely, graft com- diagnostic tests for graft compatibility. Perhaps serological
patibility-incompatibility is a combination of the tests for graft compatibility may be developed in the
auxin-toxin interactions of Figure 11–43 and/or some future, to complement those currently used in disease
chemical recognition response. To date, we have little diagnostic kits of plant pathogens.
understanding of the molecular chain of events that The introduction of new Prunus rootstock can be
occurs during wounding (180) and graft union forma- difficult (and very costly!) because incompatibility can
tion, or how those chains of events vary between com- occur some years after grafting. The composite tree can
patible-incompatible graft partners. In Douglas-fir, grow “normally” for years, and then a breakdown occurs
graft incompatibility is apparently controlled by multi- at the graft union area. It is now known that with incom-
ple genes with additive effects (36). patible apricot-plum (Prunus) grafts, some callus differen-
tiation into cambium and vascular tissue does occur;
however, a large portion of the callus never differentiates
Predicting Incompatible Combinations (Fig. 11–40) (48). Early detection of graft incompatibility
Accurately predicting whether or not the components of in fruit trees is greatly facilitated since this process can be
the proposed scion-stock combination are compatible detected histologically Magnetic Resonance
would be tremendously valuable. An electrophoresis test was within weeks after Imaging (MRI)
used for testing cambial peroxidase banding patterns of the grafting (48). A diagnostic imaging tech-
scion and rootstock of chestnut, oak, and maple (138, Magnetic reso- nique that can be used for
140–145). Peroxidases mediate lignin production. nance imaging detecting vascular conti-
Increased peroxidase activity occurs in incompatible (MRI) can be used to nuity in the callus bridge.
Figure 11–43
A model to explain the development of a
compatible graft union. The stages are
adhesion of the scion and rootstock,
proliferation of callus cells to form the callus
bridge, and vascular differentiation across
the graft interface. The outer callus cells are
from the periderm and outer cortex.
The pressure exerted on the graft is from
the physical contact of the scion to the
rootstock—and the development of a
suberized periderm. Auxin is a potential
morphogen, enhancing vascular dediff-
erentiation. In this model, incompatibility is
not caused by specific cellular recognition
events between the graft partners. Rather,
incompatibility may occur when a toxin,
such as hydrocyanic acid (HCN) or
benzaldehyde, counteracts naturally
occurring morphogens (e.g., auxin), thus
inhibiting or degenerating vascular tissues
in the graft union (106).
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 450
Figure 11–44
The graft interface of an incompatible graft
between Sedum telephoides and Solanum
pennellii at eight days after grafting. Lethal
cellular senescence in Sedum has resulted in
the formation of a necrotic layer of collapsed
cells that separates the two graft partners.
×5,000. Courtesy R. Moore and D. B. Walker (101).
reactions. In practice, it may be difficult to separate with dwarfing rootstocks, and delay in fruiting with
which influencing factor is dominant in any given graft vigorous rootstocks. Apple rootstocks are used prima-
combination growing in a particular environment. rily for reducing tree size and for increasing precocity
Long-term results depend on the rootstock-scion and yield efficiency.
combination, environment (climate, edaphic factors Besides being more precocious, intensive plant-
such as soil), propagation, and production manage- ings of small trees resulting from dwarfing rootstock
ment, which affects yield, quality, plant form, and intercept more light and have less internal shading,
ornamental characteristics (if applicable), and by exten- which is related to greater dry matter production and
sion, the economics of production. fruit yield. The higher ratio of fruit weight to trunk and
branch weight (partitioning of photosynthate to fruit
Effects of the Rootstock rather than wood formation) may also contribute to
on the Scion Cultivar higher yield efficiencies for trees growing on dwarfing
rootstock than more vigorous clonal and seedling root-
Size and Growth Habit Size control, sometimes
stock (121, 160).
accompanied by change in tree shape, is one of the
Vigorous, strongly growing rootstocks, in some
most significant rootstock effects. Rootstock selection
cases, result in a larger plant that produces a bigger crop
in apple has produced a complete range of tree sizes—
(per individual tree) over many years. On the other
from dwarfing to very vigorous—by grafting the same
hand, trees on dwarfing rootstocks are more fruitful,
scion cultivar to different rootstocks (Fig. 11–10).
and if closely planted, produce higher yields per hectare
That specific rootstocks can be used to influence
(acre). The producer’s cash flow and return on invest-
the size of trees has been known since ancient times.
ment are much improved because an apple crop on
Theophrastus—and later the Roman horticulturists—
dwarfing rootstock produces more fruit earlier.
used dwarfing apple rootstocks that could be easily
Furthermore, the management costs of harvesting,
propagated. The name “Paradise,” which refers to a
pruning, spraying, and general maintenance are much
Persian (Iranian) park or garden—pairidaeza—was
greater on large trees.
applied to dwarfing apple rootstocks about the end of
the 15th Century.
Size, Quality, and Maturity of Fruit There is consider-
A wide assortment of size-controlling rootstocks
able variation among plant species in regard to the effect
has now been developed for certain of the major tree fruit
of the rootstock on fruit characteristics of the scion cul-
crops. Most notable is the series of clonally propagated
tivar. However, in a grafted tree there is no transmission of
apple rootstocks collected and developed at the East
fruit traits characteristic of the rootstock to the fruit pro-
Malling Research Station in England, beginning in 1912.
duced by the grafted scion. For example, quince, com-
These apple rootstocks were classified into four groups,
monly used as a dwarfing pear rootstock, has fruits with
according primarily to the degree of vigor imparted to the
a pronounced tart and astringent flavor, yet this flavor
scion cultivar: dwarfing, semi-dwarfing, vigorous, and
does not appear in the pear fruits. The peach is often
very vigorous—same size as seedling rootstock
used as a rootstock for apricot, yet apricot fruits do not
(Fig. 11–10). Similarly, the size-controlling effects of the
have any characteristics of peach fruits.
rootstock on sweet cherry (Prunus avium) scion cultivars
Although there is no transfer of fruit characteristics
has been known since the early part of the 18th Century.
between the rootstock and the scion, certain rootstocks
Mazzard (P. avium) seedling rootstocks produce large,
can affect fruit quality of the scion cultivar. A good exam-
vigorous, long-lived trees, whereas P. mahaleb seedlings, as
ple of this is the “black-end” defect of pears. ‘Bartlett,’
a rootstock, tend to produce smaller trees that do not live
‘Anjou,’ and some other pear cultivars on several different
as long. However, individual seedlings of these species,
rootstocks often produce fruits that are abnormal at the
when propagated asexually and maintained as clones, can
calyx end. While the fruit quality and yield of tomatoes
produce different, distinct rootstock effects. Rootstock
and cucurbits is generally enhanced with the correct
effects on tree size and vigor are recognized also in citrus,
stock-scion combination, sometimes melon fruit quality
pear, peach, olive and other species. A discussion of spe-
is impaired when grafted on disease resistant Cucurbita
cific rootstocks for the various fruit and nut crops is given
rootstock (39). Rootstocks of chili peppers (Capsicum
in Chapter 19.
annuum) can increase the level of capsaicin, which influ-
Fruiting Fruiting precocity, fruit bud formation, fruit ences the “hotness” of peppers (185).
set, and yield of a tree can be influenced by the root- In citrus, striking effects of the rootstock appear
stock used. In general, fruiting precocity is associated in fruit characteristics of the scion cultivar (23). If sour
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 452
Disease and Pest Resistance. Some rootstocks are interstock, even in rootstock types that normally do not
more tolerant to adverse soil pests, such as nematodes sucker freely. Today, apple interstocks are rarely used
(Meloidogyne spp.), than others; for example, except in China (G. Fazio, personal communication).
‘Nemaguard’ peach rootstock. The growth of the scion
cultivar is subsequently enhanced by the rootstock’s
Possible Mechanisms for the Effects
ability to withstand these soil pests. Grape cultivars sus-
of Rootstock on Scion and Scion
ceptible to the insect pest, phylloxera (Dacylosphaera
vitifoliae), are grafted onto resistant rootstocks. Many on Rootstock
cucurbits and solonaceous crops are grafted for While many of the effects of rootstock-scion relations are
enhanced disease resistance and tolerance of abiotic known, the fundamental mechanisms of control, partic-
stress (Figs. 11–3 and 11–4) (34, 39, 82, 86, 131). ularly on the molecular basis, are not well understood.
Grafting with disease-resistant rootstock also offers new A better understanding of the mechanisms controlling
IPM management strategies for organic vegetable pro- growth and development in grafted plants would speed
duction (131). up the design, development, and commercialization of
new composite plant systems. By understanding these
Effect of the Scion Cultivar mechanisms, breeders could better predict the growth
on the Rootstock responses of new potential graft partners (while they are
still on the “drawing board”) and develop more efficient
Although there is a tendency to attribute all cases of
screening tests—rather than relying on cumbersome trial
dwarfing or invigoration of a grafted plant to the root-
and error processes that may take up to 10 years or more
stock, the effect of the scion on the behavior of the com-
in evaluating grafted, woody perennial plants.
posite plant may be as important as that of the rootstock.
Without question, the nature of the rootstock-scion
Effect of the Scion on the Vigor and Development of the relationship is very complex and differs among genetically
Rootstock Scion vigor can have a major effect on root- different combinations. Furthermore, in a composite
stock growth, just as rootstocks can affect scion growth. plant system, size control, plant form, flowering, fruiting,
If a strongly growing scion cultivar is grafted on a weak disease resistance, flood tolerance, etc. are not controlled
rootstock, the growth of the rootstock will be stimulated by the same genes or physiological/morphological mech-
so as to become larger than it would have been if left anisms. Theories advanced as possible explanations for
ungrafted. Conversely, if a weakly growing scion cultivar the interaction between the rootstock and scion include:
is grafted onto a vigorous rootstock, the growth of the (a) anatomical factors, (b) nutritional and carbohy-
rootstock will be lessened from what it might have been drate levels, (c) absorption and translocation of nutri-
if left ungrafted. In citrus when the scion cultivar is less ents and water, (d) phytohormones and correlative
vigorous than the rootstock cultivar, it is the scion culti- effects, and (e) other physiological factors.
var rather than the rootstock that determines the rate of
growth and ultimate size of the tree (66). Anatomical Factors The roots and stems of dwarfing
apple rootstocks, which can reduce vegetative growth and
increase flowering, are characterized by several anatomical
Effect of Interstock on Scion features. These include: (a) a high ratio of bark (periderm,
and Rootstock cortex, and phloem tissue) to wood (xylem tissue); (b) a
The ability of certain dwarfing rootstock clones, large proportionate volume occupied by living cells (axial
inserted as an interstock between a vigorous top and parenchyma and ray parenchyma cells) relative to func-
vigorous root, to produce dwarfed and early bearing tionally dead xylem cells (vessels and fibers); and (c) fewer
fruit trees has been used for centuries to propagate and smaller xylem vessels (13, 14, 92, 153).
dwarfed trees. The degree of size control induced Much of the functional wood tissue of roots of
in apples by various dwarfing rootstock is shown in dwarfing apple stocks is composed of living cells, whereas
Figure 11–10. Dwarfing of apple trees by the use of a in nondwarfing, vigorous rootstocks, the wood consists of
‘Malling 9’ as an interstock was a common commercial a relatively large amount of lignified tissue without living
practice for many years (Fig. 11–11). This dwarfing cell contents (i.e., a larger vessel/tracheid system for more
method had the advantage of allowing the use of well- efficient water transport). At the graft interface between
anchored, vigorous rootstock rather than a brittle, the scion bud and dwarfing apple rootstock, xylem vessels
poorly anchored dwarfing clone. Sometimes excessive with smaller than normal diameter are formed, whereas
suckering from the roots occurred due to the dwarfing semi-dwarfing rootstock produces normal xylem after a
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 454
brief interruption (157). It has been proposed that failure harvested, the insoluble root starch supply is greater, but
of auxin to cross the bud-union interface in the case of soluble sucrose and sorbitol are less in vigorous root-
the dwarfing rootstock leads to reduced rootstock xylem stock compared with dwarfing rootstock.
formation, and hence a reduced supply of water and min- It appears that apple rootstock does not influence
erals to the scion, thus causing the dwarfing effect (87). mineral nutrition at the site of flower formation (65).
Defects in the graft union that cause a partial discontinu- Most likely, rootstock effects on flowering are due to
ity of the vascular tissues may in part explain the marked internal control mechanisms that affect the proportion
depletion of solutes, nutrients, and cytokinins (produced of spurs that become floral (64).
from root apices) in the sap contents of dwarfing inter- To summarize, dwarf apple rootstocks do affect
stocks and rootstocks (73). precocity and flowering, in part, because of differences
Conversely, with kiwifruit, the roots of flower- in carbohydrate metabolism and the greater carbon
promoting rootstock tend to have more and larger xylem partitioning to the reproductive areas. The contribut-
vessels, more crystalline idioblasts, and more starch grains ing influence of hormones, which also affects carbon
(173, 174). Most likely the greater water supply from the partitioning and flowering, is discussed below.
rootstock to the scion in early spring determines the
abundance of flower production of the kiwifruit scions. Absorption and Translocation of Nutrients and Water
Morphologically, dwarfing rootstock have fewer Apple rootstocks affect Ca, Mg, Mn, and B uptake, but
coarse roots (diameter greater than 2 mm) and fewer there is no apparent direct relationship of mineral sta-
fine roots (diameter less than 2 mm) than more vigor- tus with rootstock vigor, productivity, or spur charac-
ous apple rootstocks (5, 6). There is not always a clear teristics (65).
relationship between root length growth and size con- Rootstocks do differ in their ability to absorb and
trol characteristics of dwarfing versus vigorous root- translocate P (74), but a direct role of phosphorus at
stock. However, there are fewer active root tips in the site of flower formation induced by rootstock seems
dwarfing than vigorous apple rootstock (51). The unlikely (65). In a study of the translocation of radioac-
roots and shoots of vigorous apple rootstocks also have tive phosphorus (32P) and calcium (45Ca) from the
a longer growing season than dwarfing rootstock (78). roots to the tops of 1-year ‘McIntosh’ apple trees grown
Nutritional and Carbohydrate Levels Dwarfing root- in solution culture, it was shown that more than three
stock of apple tends to partition a greater proportion of times as much of both elements was found in the scion
carbon to reproductive areas (spurs, spur leaves, fruit) and top when vigorous rootstock was used in comparison
less to the tree branch and frame dry weight, compared with the dwarfing rootstock (29). This may indicate a
with nondwarfing rootstocks (160). The greater water and superior ability of the vigorous rootstock to absorb and
nutrient uptake of the vigorous rootstock contributes translocate mineral nutrients to the scion in compari-
to the production son with the dwarfing rootstock. Or it may only mean
spurs The principal fruiting of new vegetative that roots of the dwarfing rootstock, with their higher
unit in apple, which may be growth, which is a percentage of living tissue, formed a greater “sink” for
classified as short shoots. competing sink these materials, retaining them in the roots.
The terminal bud of a spur with reproductive Interstocks of such dwarfing apple clones as
may be either vegetative, growth. ‘Malling 9’ will cause a certain amount of dwarfing,
containing only leaves, or The root- suggesting reduced translocation due to partial block-
reproductive. Reproductive stock affects the age at the graft unions or to a reduction in movement
buds of apple are mixed partitioning of the of water or nutrient materials (or both) through the
buds that produce both dry matter between interstock piece. Differences among rootstocks in water
flowers and leaves. above- and below- translocation have been demonstrated with a steady-
competing sink The ground tree com- state, heat-balance technique that accurately measures
competition of two ponents. Vigorous xylem sap flow rate and sap flow accumulation over
independent growth rootstocks accumu- time. Under nonstress conditions, sap flow was greater
processes (such as flowering late more dry mat- in ‘Granny Smith’ apple scions grafted to very vigorous
and adventitious root ter in the shoot and seedling (standard) rootstock, while sap flow was simi-
formation) for the same root system than lar between the dwarfing and semi-dwarfing rootstock
limited metabolic resources dwarfing stock (6, (70). Moisture stress affects the sap flow of the vigorous
(e.g., carbohydrates, 161). At the time seedling rootstock the least and reduced sap flow on the
proteins). apples are being dwarfing rootstock the most.
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 455
Sweet cherries grafted on dwarfing rootstock have into the root system, where they affect root growth.
smaller and fewer xylem vessels in the scion and graft Cytokinins are produced predominantly from root
union, and irregular vessel orientations in the vascular apices, and are translocated primarily through the
tissue compared to non-dwarfing rootstock; this differ- xylem, where they can influence physiological responses
ence could contribute to greater hydraulic resistance in and growth in the scion.
the graft union, resulting in reduced scion growth Of the phytohormones, auxin plays one of the
(dwarfing) (116). With peach trees grafted on rootstocks most important roles in dwarfing rootstock control of
with differing size-controlling potentials, the higher root apple scion growth (75). The dwarfing effect may be
resistance (reduced sap flow) plays a central role in the explained by reduced auxin transport into the graft
dwarfing mechanism induced by size-controlling root- union of the dwarf rootstock (87); this could alter the
stock (semi-dwarfing). Interestingly, the root system hormonal balance between shoots and roots, and
accounted for the majority of resistance of water flow account for the reduced vegetative growth and vigor of
through the tree and had no effect on hydraulic conduc- the scion. Auxin affects vascular differentiation, and is
tance through the scion or graft union (12). important for stimulating cambial activity and xylem
Conversely, with olive trees (Olea europaea L.), development (1) in the graft union area and the vascular
while there was lower hydraulic conductance in dwarf- system of the grafting partners. Dwarfing yields greater
ing than vigorous rootstock during the first several reduction in cambial activity and xylem formation in the
months, but after 1 year hydraulic conductivity was the graft union than vigorous rootstock (158) because of the
same between dwarfing or vigorous rootstocks (55). dwarfing’s reduced capacity to support polar auxin trans-
In summary, as long as mineral elements are not port (not auxin uptake into cells), and a reduced capacity
limiting, the greater uptake of P and Ca by the more for auxin efflux from transporting cells (158). Since
vigorous rootstock does not adequately account for auxin is known to stimulate its own transport (58), lower
dwarfing effects (29). While rootstocks can influence endogenous auxin levels in the dwarfing rootstock may
leaf mineral nutrition, results have been inconsistent limit its capacity to support polar auxin transport. A
(65). In general, sap flow is greatest in vigorous and least chain of events is set off with less auxin being trans-
in dwarfing rootstock. While differences in sap flow ported, which leads to reduced cambial activity and sub-
may be attributed to differences in root characteristics, sequently reduced xylem formation. Reduced xylem for-
xylem anatomy or other features of the hydraulic archi- mation limits conduction in the dwarf rootstock, which
tecture from the roots to the graft union, or the union concurs with the reports on lower xylem sap flow (70).
itself, the primary influence probably lies more in the There is evidence for greater auxin accumulation
nature of the growth characteristics of such rootstocks. in the scion of dwarfing apple understock. With apple,
hydraulic conductivity of the graft tissue was lower for
Phytohormones and Correlative Effects Plants main- grafted trees on dwarfing rootstocks, compared to
tain a constant root/shoot ratio, and any attempt to alter semi-vigorous rootstocks. The amount of functional
this ratio results in the plant redirecting its growth pat- xylem tissue in the graft union and scion initially
tern until the ratio is reestablished. This also applies to increased with rootstock vigor (7). However, as the
grafted plants and plants grafted tree aged, any differences in sap flow become mar-
correlative effects
transplanted into a land- ginal. The dwarfing tree compensated for hydraulic
The influence of one
scape site or orchard. limitations imposed by the graft tissue and abnormal
organ over another, due
Producing a composite xylem anatomy (compared to more vigorous rootstock)
to phytohormones (e.g.,
plant by grafting onto a by initially reducing its transpiring leaf area, and pro-
high ABA produced in
dwarfing rootstock is an ducing a smaller canopy (smaller tree). As the dwarfed
the root tips of dwarfing
alteration in the normal tree aged, the cross-sectional area of the graft union
apple rootstock reduces
growth pattern (87). increased (7), brought about by greater auxin accumu-
the vegetative growth
Growth in the composite lation (reduced transport) in the graft tissue of the
of the scion).
plant will be redirected dwarfing rootstock, which led to increased xylem devel-
until equilibrium is reached between the rootstock-scion opment later, as the dwarf tree aged.
system. Intimately involved in redirecting plant growth Auxin can indirectly affect cytokinin production.
are the correlative effects of root (rootstock)/shoot Reduced auxin transport leads to a smaller root system
(scion) systems, mediated by phytohormones. Auxins, in the dwarf rootstock that produces less cytokinin,
which are produced predominantly in the shoot system, and/or the root metabolism is sufficiently altered to
are basipetally translocated through the phloem and affect cytokinin synthesis. Subsequently, there is less
M11_DAVI4493_08_SE_C11.qxd 8/26/10 7:33 PM Page 456
cytokinin translocated upward from the roots to the concluded there was little evidence to support a role for
shoots and reduced top growth occurs; hence, the gibberellin in vigorous, compared to dwarfing rootstock
dwarfing effect. This correlative effect is mediated by (87, 134, 135). However, in other studies, dwarfing
auxin and cytokinin as growth in the composite plant is (M9) interstock labeled GA3 was lower, and glycosyl
redirected and equilibrium is reached between the conjugated GA3 (inactive GA3 form) was higher com-
dwarf rootstock/scion system. pared to nondwarfing (MM115) interstocks (130).
Abscisic acid (ABA) and gibberellic acid (GA) However, a problem with hormonal studies is that the
may also play a role in the correlative effects of dwarf- composition of the xylem sap often has very little resem-
ing rootstock. Root apices are an important site of blance to that flowing through the intact, transpiring
ABA synthesis. The dwarfing ‘Malling 9’ apple root- trees. Hormone and ion concentrations in osmotically
stock contains lower amounts of growth-promoting exuding sap do not always reflect the condition of the
materials—but more growth inhibitors—than does intact plant (5). For instance, slow-flowing sap concen-
the very vigorous ‘Malling 16’ rootstock (88). ABA trates solutes faster than fast-flowing sap diluted solutes.
levels are also reported to be higher in dwarfing root- Apparently, xylem-borne substances are not delivered in
stock (184), and in the stems of dwarfed apple trees, proportion to sap flow, suggesting that differences in
than in more vigorous ones (135). tree transpiration or leaf area have considerable influ-
There are higher ratios of ABA:IAA (auxin) in ence on signal molecule concentration and delivery (5).
dwarfing than vigorous apple rootstock, a finding con- In summary, with apple, auxin is directly involved
firmed using gas chromatography-mass spectropho- in dwarfing rootstock effects, and cytokinins (which are
tometer selective ion affected by auxin-mediated root growth and subse-
marker A morpho- monitoring techniques quent cytokinin biosynthesis) are either directly or
logical, biochemical, (78). Higher ABA:IAA indirectly involved in plant size control. There is a
genetic indication of a ratios may lead to strong case for ABA-mediated dwarfing effects, while
trait (e.g., higher ABA in greater differentiation there are conflicting reports on the role of GA. Most
shoot bark of dwarfing of phloem and related likely, there is an interaction of factors affecting dwarf-
compared with a tissues in dwarfing root- ing phenomena such as phytohormones, anatomical
vigorous apple stocks, which could factors, nutrition and carbohydrate levels, sap flow, and
rootstock). explain why dwarfing translocation of carbohydrates across the graft union.
rootstocks have higher
bark (periderm, cortex, phloem, vascular cambium) to Other Physiological Factors A wide range of physio-
wood (xylem) ratios than vigorous rootstocks. The logical characteristics have been found to affect root-
higher concentration of ABA in shoot bark of dwarfing stocks, scions, and their resulting interactions (87, 127,
compared with vigorous rootstock is a potentially useful 162). For example, rootstocks have been found to influ-
marker in selecting for dwarfing apple rootstock (78). ence transpiration rate and crop water-use efficiency in
There are conflicting reports that higher GA is peach; leaf conductance and osmotic potential in apple;
found in more vigorous rootstock. Earlier reports and midday leaf water potential in citrus, peach, and
apple trees. Rootstock-scion combinations can also Cytokinins are known to promote photosynthesis, and
influence net photosynthesis and growth characteristics root-produced ABA—translocated in xylem sap—can
of grafted Prunus species under droughted conditions reduce stomatal conductance and photosynthetic rates
(127). The greater tolerance to flooding found in in the shoot system.
selected rootstock of Prunus (124) and fir (Abies) is More needs to be done with the molecular basis
probably due to physiological and/or morphological of rootstock-scion relations. It is possible that certain
mechanisms (45) that allow selected rootstock to handle genes are being turned on and off and/or that genetic
anaerobic conditions better than other rootstock. information may be transmitted between the graft part-
Net photosynthesis of leaves tends to be higher ners of the composite plant (115). Epigenetic changes
with apple scions on vigorous rootstock than on dwarf- occur in grafting with the speeding up of maturation
ing rootstock (148). But photosynthetic rates cannot on grafted versus seedling-grown plants (see discussion
be used to explain differences in yield and yield effi- on epigenetic changes in Chapter 16). Conversely,
ciencies induced by the rootstock. Part of this complex- micropropagated dwarfing apple rootstocks that are
ity is because the presence of fruit increases leaf net grafted can have more juvenile-like characteristics,
photosynthesis by some unknown mechanism (148). which delays bearing and fruit cropping of trees (76).
DISCUSSION ITEMS
1. What have been some historical reasons for graft- 8. Why is there potentially more rapid graft union
ing compared to other propagation methods? development and frequently a higher percentage of
2. Compare budding and grafting. “takes” in chip budding compared to T-budding?
3. What are the differences between seedlings and 9. What environmental conditions are desirable dur-
clonal rootstock? What are the advantages of each ing and following grafting?
system? 10. What are the genetic limits of grafting,(i.e., when
4. Using an interstock (double working) is expensive. is grafting most likely to be successful)?
Why is it still used as a propagation technique? 11. What are the different types of graft incompatibil-
5. What are some of the ecological advantages of nat- ity, and what causes them?
ural root grafting? How can it be a disadvantage in 12. What are some techniques to help predict graft
the dissemination of diseases, such as oak wilt and incompatibility?
Dutch elm disease? 13. What are some possible mechanisms for size con-
6. What are the stages of graft union formation? trol (dwarfing) in stock-scion relations?
7. Does cellular recognition take place in grafting,
and, if so, how might that be important to graft
compatibility/incompatibility?
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