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275
CHAPTER 13
Selection methods
Part 5: Breeding clonally propagated
crops
Wolfgang Grüneberg, Robert Mwanga, Maria Andrade and Jorge Espinoza

276 Plant breeding and farmer participation
13.1 INTRODUCTION 13.2 AN OVERVIEW OF CLONALLY

The literature about participatory breeding PROPAGATED CROPS
of clonally propagated crops is very limited. What are clonally propagated crops?
This makes it difficult to write this chapter Standard textbooks list a surprisingly large
exclusively about how farmers have bred, number of crops: all important root and
how they breed—with or without support tuber crops, many forage crops, nearly all
of scientists—and how they should breed types of fruit and wooden ornamentals,
clonally propagated crops. There are many cut flowers and pot plants, as well
today clear definitions for participatory as forest trees. The definition of a clonally
plant breeding (PPB), participatory propagated crop is that the material to
variety selection (PVS) (see Chapter 9) cultivate and maintain a variety is obtained
and indigenous plant breeding (IPB; the by asexual reproduction, regardless of
selection process of farmers for more than how different the plant material used for
60 centuries). Breeding by scientists and propagation is within and between species,
economic entities has been redefined as encompassing tubers, roots, stem cuttings
formal plant breeding (FPB), consisting and corms, as well as asexually developed
of breeding carried out on-station, linked seeds (seeds developed without meiosis). It
with multi-location trials, and assisted should be remembered that if crops such as
by quantitative genetics, selection theory maize (bred as an open-pollinated or hybrid
and biotechnology (with or without crop) or beans (bred as a cross-fertilized,
recombinant DNA technology). A variety self-fertilized or hybrid crop) were to be
developed through FPB is termed a modern propagated by stem cuttings or asexually
variety (MV), in contrast to a farmer-bred developed seeds, they would be clonally
variety (FV), which is developed by IPB. propagated crops. In contrast, in breeding
Especially in clonally propagated crops, FVs clonally propagated crops, the breeding
continue to dominate crop production in techniques and methods that are usually
many developing world regions. Obviously associated with cross-fertilized and hybrid
FPB has not been so successful, because crops can be very useful. An example is
farmers decided to continue to grow FVs the selection of parents in potato, cassava
instead of adopting MVs (Friis-Hansen, and sweet potato breeding, which are
1992; Witcombe et al., 1996). Definitions recombined in open-pollinated polycross
are helpful, providing a common term for nurseries to create new genetic variation.
the same technique or method. Here we It is almost certain that techniques and
need to bear in mind two points when methods from breeding cross-fertilized
considering such definitions and the issues and hybrid crops will become much more
associated with them: important in the future of clone breeding.
• the every-day formula used to predict
the response to selection (with all its What is the general principle in
extensions to several selection steps and breeding clonally propagated crops?
traits) may well be the most useful tool It appears to be simple: to break the normal
given to breeding by statistics; and clonal propagation by a crossing step, and
• breeders have to adapt a crop to human thus develop sexual seeds and genetic vari-
needs and they must pay adequate ation from which to select new clones. All
attention to the needs of clients. propagation steps from the first to the last
Selection methods. Part 5: Breeding clonally propagated crops 277
selection step are again ‘normal’ asexual result was the domestication of pitiquiña
reproduction (Simmonds, 1979). Hence, (Solanum stenotomum), which was most
the finally selected clone is genetically iden- probably selected from S. leptophyes or
tical with the original seed plant from S. canasense. From the view-point of the
which the selected clone is derived. In other knowledge of the twenty-first century it is
words, each seed plant is a potential variety. not surprising that suddenly potato plants
Roots and tubers, fruit and tree plant spe- with larger leaves and larger tubers were
cies have been used by human since long found. Potato spontaneously changes its
before the dawn of agriculture. They have polyploidy level by unreduced gametes and
been domesticated by IPB (Simmonds, recombination. Polyploid potatoes are more
1979) and several made a substantial yield vigorous than their diploid ancestors. The
progress by FPB in some regions of the result was the domesticated of polyploid
world. However, in other regions of the andigena (S. tuberosum subsp. andigena).
world there is not much yield progress, Andigena is the ancestor of the commercial
and in these regions there appears to be a potato in long-day temperate climates—the
clear need of PPB for progress. We wish to so-called Irish potato (S. tuberosum subsp.
illustrate this by two examples: potato and tuberosum) (Hawkes, 1979, 1981). This IPB
sweet potato. of potato and introductions of FV of potato
An example of the needs and requirements into the Northern Hemisphere changed
of clonally propagated varieties can be the world both socio-economically and
found in potato (Solanum spp.). There are politically (Hobhouse, 1985).
about 200 wild potato species (Huamán Today, eight species of potato are still
and Ross, 1985). They usually contain cultivated in the Andes, variously diploid,
glycoalkaloids, which give tubers a bitter triploid, tetraploid and pentaploid:
taste and which are toxic when consumed (i) cultivated diploid potatoes are
in large quantities (Zitnak and Filadelfi, pitiquiña (S. stenotomum), its close
1985). It is nearly certain that 100 to 130 relatives phureja (S. phureja) and
centuries ago indigenous knowledge in limeña (S. goniocalyx), and ajanhuiri
the Andes and along the Pacific coast of (S. ajanhuiri), which evolved from
South America was those sites where it was interspecific recombination of diploid
possible to collect wild potato tubers where pitiquiña and the diploid wild potato
species and mutants were growing that had species S. megistacrolobum;
low alkaloid content. Although these tubers (ii) cultivated triploid potatoes are
were very small, the man, or more probably chaucha (Solanum × chaucha), a
a woman, made life much easier by growing hybrid between diploid pitiquiña
and maintaining desirable types by cloning and tetraploid andigena, and rucki
close to their homes. This happened more (Solanum × juzepczukii), a hybrid
than 8 000 years ago, and most likely between diploid pitiquiña and the
independently at several places (Ugent, tetraploid wild potato species
Pozorski and Pozorski, 1982; Ugent, S. acaule;
Dillehay and Ramirez, 1987). Those types (iii) cultivated tetraploid potatoes are
were preferred that were easier to maintain, andigena and Irish potato; and finally
easier to harvest (shorter stolons) and had (iv) the cultivated pentaploid potato
larger tubers compared to other types. The is a hybrid species (Solanum ×
curtilobum), which evolved between important as taste, flavour and nutrient

tetraploid andigena and triploid rucki, content, namely the importance of
and unfortunately is also called rucki adaptation of a crop and its varieties to
(Hawkes, 1981; NRC, 1989). the local environment. They who know
The andigena is the best known potato in the Andes also know that is unrealistic
the Andes (with about 2 500 known FVs). to breed a widely adapted potato variety
It is cultivated in tropical mid-elevation for this region of the world. Temperature,
valleys and mountainsides. The second rainfall, soil conditions (including salinity
most important potato is phureja, which is and drought) and pest and disease pressures
cultivated on the warm and moist eastern change from microclimate to microclimate
slopes of the Andes (with about 500 known from sea level at the Pacific coast up to 3 500
FVs), followed by limeña, ajanhuri and to 4 500 masl in the Andean highlands (mid-
rucki. Limeña or papa amarilla is grown elevation valleys and plateaus), and again
in the temperate areas of the Andes and down into the warm tropics, where the
still achieves high market prices due to its Andes meet the Amazon. Breeding potatoes
taste and flavour. Ajanhuri and rucki are in this region of the world was and can only
the most frost resistant cultivated potato be successful by decentralization and with
species and cultivated up 4 200 masl. The farmer participation (Johns and Keen, 1986;
former is used as an insurance crop in cases Gabriel and Torrez, 2000). Admittedly this
where andigena fails due to unpredictable is an extreme example, but such situations
hail and frost (some ajanhuri varieties are can be found in less extreme form in nearly
bitter and must be processed). The latter all regions of the world. In the Southern and
are usually only eaten after having been Northern Hemispheres, potatoes generally
processed into chuño, the famous storable must be day-neutral; in South-west and
food product of the Incas. Many of these Central Asia, potatoes must be very quick to
potatoes have clearly better taste and flavour mature, with a short crop duration of 80 to
compared with what is considered potato in 90 days; in Europe and Northern America,
the Northern Hemisphere (Huamán 1983; more than 30 quality characteristics combine
NRC, 1989; De Haan, 2009). However, taste to determine tuber quality for market needs;
is a variable characteristic; it changes from and finally in the UK, a potato variety must
person to person, from family to family, and be white fleshed, whereas in Germany it
from society to society. Moreover, many of must be yellow fleshed, otherwise it is
these IPB potatoes are clearly superior in not eaten (CIP, 1984; Levy, 1984; Tarn
protein and micronutrient concentration et al., 1992). An additional major factor
in their tubers (pro-vitamin A, calcium, for adaptation and acceptance of potato
magnesium, iron and zinc) compared with varieties is their tolerance and resistance to
MVs (Ochoa, 1990; Morris et al., 2004; diseases and pest. In all temperate and moist
Burgos et al., 2008), and are useful as genetic climates, potato farmers have to fear Late
resources or directly as FVs to alleviate blight (Phytophthora infestans), which is
malnutrition in the mountain regions of not important at temperature above 25°C,
the world. but then Early blight (Alternaria solani)
The potato and the Andes were chosen takes over. In tropical lowlands, the farmer
as an example to give an impression of has to fear Bacterial wilt (Pseudomonas
an aspect of breeding that is as least as solanacearum), and in all warm dry regions
the potato crop can be lost because of high DM, low-sweet or bland type, which
Colorado beetle (Leptinotaras decemlineata) has a dry mouth feeling. The first type, also
(CIP, 1977, 1980; Radcliffe, 1982; Rich, called the dessert type, has extremely high
1983). pro-vitamin A concentrations (Huang,
A simpler example for the needs and Tanudjaja and Lum, 1999) and a 50 g piece
requirements of clonally propagated varie- of fresh storage roots can meet the daily
ties can be found in sweet potato (Ipomoea requirements of a pre-schooler (Low et
batatas). Sweet potato was domesticated in al., 2007). Moreover, sweet potatoes with
the Americas more or less during the same high pro-vitamin A concentrations have
prehistoric period as the potato (O’Brien, high protein and mineral concentrations
1972). The evolution of sweet potato was (Grüneberg, unpublished). In the United
not as complex and diverse as that of States of America, the dessert type is
potato. There are about 500 Ipomoea spe- generally the desired sweet potato to
cies, but only the I. batatas species was meet market and consumer needs. In the
domesticated (Austin and Huamán, 1996). Caribbean, low DM orange-fleshed sweet
Again polyploidy was important. Sweet potatoes (OFSP) are consumed, but as
potato is hexaploid and its closest relative a staple, so a dryer mouth feel and less
is I. trifida (di- and tetraploid). It is certain sweet flavour is preferred. These white-
that sweet potato contains the I. trifida or yellow-fleshed varieties are known as
genome, but obviously it is not simply a bonitos or ricos (Baynes, 1972). Along the
multiple copy. Two-thirds of the sweet Pacific coast of South America we observed
potato genome corresponds to the I. trifida that sweet potatoes are mainly pale orange
genome and one-third to an ancestor very fleshed and less sweet. However, locally,
closely related to I. trifida (Shiotani and white- and purple-fleshed sweet potatoes
Kawase, 1989). Within diploid I. trifida are consumed, which clearly have different
accessions (seed families) it is also possible taste, texture and flavour compared to
to find plants that form small storage roots OFSP. In Brazil, the sweet potato storage
(Daniel Reynoso, pers. comm.). However, roots must clearly have a high DM
sweet potato has been found in the ruins concentration (28 to 30 percent DM), and
of the so-far oldest city in the Americas, usually this is a white-fleshed sweet potato;
Caral on the Pacific coast of central Peru however, locally, high DM OFSP can be
(Solis, 2004), and the crop reached Pacific found (Amauri Buso, pers. comm.).
Polynesia and parts of South-East Asia The taste preferences in sub-Saharan
(naturally or by early seafarers) before Africa are similar to those in Brazil, per-
Columbus. It was primarily the Portuguese haps because the Portuguese introduced the
that introduced it into Europe, Africa, sweet potato into Africa. All FVs are nearly
South Asia and East and South-East Asia exclusively white- or yellow-fleshed and
(Yen, 1976). have high DM concentrations; however, a
Although the taste of sweet potato in FVs few pale- to medium-orange-fleshed FVs
and MVs differ tremendously, two major can be found with high DM concentrations
types can be distinguished: (i) the orange- (Tumwegamire, unpublished). These local
fleshed, moist, low dry matter (DM) and OFSP FVs are very promising for allevia-
sweet type, which has a soft mouth feeling; tion of vitamin A deficiency in sub-Saharan
and (ii) the white- or pale-yellow-fleshed, Africa (e.g. FVs such as ‘Ejumula’, ‘Carrot
C’, ‘Carrot Dar es Salaam’ or ‘Zambezi’). farmers have not adopted MVs (e.g. cv.
In eastern Africa, storage root DM con- INA100, which is a high yielding OFSP
tents must be >30 percent (Mwanga et al., and fits consumer needs very well), because
2003). In southern Africa, storage DM of insufficient SPVD tolerance. Farmers
concentration between 26 and 29 percent became disappointed with new MVs and
are accepted (Laurie Sunette, pers. comm.), after a few growing seasons returned to FVs
whereas in West Africa, sweet potato must such as cv. Jonathan and cv. Huambachero.
be non-sweet, very high in DM concentra- OFSPs from the Americas with elevated DM
tions (between 30 and 35 percent DM) and (e.g. cv. Jonathan) are partially successful in
with a texture and flavour tentatively simi- southern Africa, and in south-west and
lar to yam (Dioscorea spp.) (IITA, 1981). central Asia, provided that weevil pressure is
In contrast, in India, where sweet potato not extreme. Weevil damage is associated with
consumption has been very low in the past, drought-prone regions (Central and South
today people prefer sweet potatoes with America, sub-Saharan Africa and south-
high DM, high sugar content, dark orange west and central Asia); however, weevil
flesh and a storage root shape that is cylin- species differ: Cylas formicarius in all parts
drical but tapering at both ends (Sreekanth of the tropics, C. puncticollis additionally
Attaluri, pers. comm.). in Africa, and Euscepes postfasciatus in the
In addition to regional and local West Indies. On-station and farmers’ field
preferences for storage-root colour, DM, experiments show that there are significant
texture and taste, the acceptability of sweet differences in weevil tolerance among sweet
potato varieties is mainly determined by pest potato genotypes (Hahn and Leuschner,
and disease pressures. However, the number 1981), but this tolerance appears to be less
of pest and diseases in sweet potato are pronounced or inexistent on-farm. At the
considerable lower than in potato. Generally, same time, farmers in drought-prone regions
sweet potato varieties must have a certain of Malawi want sweet potatoes in which
degree of tolerance to Sweet potato virus storage roots are formed deep in the soil
disease (SPVD). The disease occurs after and which are clearly tapering at the top,
infection by two viruses: the Sweet potato because they associate this with less weevil
feathery mottle virus (SPFMV) and the Sweet damage (Ibrahim Benesi, pers. comm.).
potato chlorotic stunt virus (SPCSV). The Moreover, latex in the storage root skin has
SPCSV is the more problematic component been associated with less weevil damage by
of SPVD, because yield losses due to SPFMV, farmers, and varieties like Santo Amaro from
in the absence of SPCSV co-infection, are Brazil clearly have considerably less weevil
low and SPFMV resistance in sweet potato damage than other sweet potato varieties
breaks after the plant is infected by SPCSV (Rafael Vasquez Martinez, pers. comm.).
(Gibson et al., 1998; Karyeija et al., 2000). The International Potato Center (CIP)
SPVD often causes serious yield losses in is promoting OFSPs to alleviate vitamin A
high-virus-pressure zones of sub-Saharan deficiency in the world (Low et al., 2007;
Africa, and American OFSPs have failed in Pfeiffer and McClafferty, 2007). However,
many regions of sub-Saharan Africa due to introductions from the Americas failed in
insufficient SPVD tolerance. Although the the high-SPVD-pressure zone of eastern
virus pressure of SPVD along the Pacific Africa (as did the FV Jonathan). To a
coast of South America is not extreme, certain extent this was associated with the
storage root flesh colour and taste. At up regional platforms for sweet potato
the same time, local African OFSP FVs, breeding in eastern, southern and West
such as Ejumula, Carrot C, Carrot Dar Africa, with a focus on dual purpose OFSP
es Salaam and Zambezi, and locally-bred (human consumption and animal feed),
OFSP MVs, such as NASPOT5 (Mwanga drought tolerant OFSP, and non-sweet high
et al., 2003), have been adopted after DM OFSP. PPB has so far mainly been a
awareness campaigns on the vitamin A research component in the organization of
deficiency problem (Regina Kapinga, pers. sweet potato breeding.
comm.). For this reason, CIP puts emphasis There are strong indications that PPB
on decentralized sweet potato breeding, in early selection steps of the sweet potato
and has recently started to recommend breeding process increases the efficiency
incorporation of at least one participatory and minimizes the risk of making wrong
selection step in the breeding process. selection decisions. In contrast to PPB, PVS
In sweet potato breeding for human is tentatively a form of on-farm evaluation
consumption, decentralization is (in the frame of a larger number of multi-
characterized by a general overall goal: that location trials) and cannot be as efficient
of developing more OFSP varieties that as PPB, because there is considerably less
meet local needs and consumer preferences, genetic variation, and, for highly heritable
to alleviate hunger and malnutrition and traits, there is nearly no genetic variation
to improve public health. The emphasis is at later breeding stages among clones. Not
on organizing OFSP breeding in eastern surprisingly, akin to the role of IPB in
and southern Africa, with national OFSP potato crop evolution, it has been shown
breeding programmes starting recently in that farmers have the ability to manage
West Africa (Ghana and Nigeria) and south- selection stages in sweet potato (Gibson
west and central Asia (India, Bangladesh and et al., 2008). This is consistent with
Sri Lanka). Breeding is almost exclusively results for potato (Gabriel and Torrez,
carried out by national agricultural 2000) and cassava (Manu-Aduening et al.,
research system (NARS) breeders and on 2006). Farmer selections are mainly made
NARS breeding stations, with currently by visual screening. This includes quality
12 NARS and two sweet potato breeders characteristics, diseases and pests, as well as
from the CGIAR system involved. NARS the growth type, which is to a certain extent
breeders are provided with funds for associated with drought adaptation in sweet
parental recombination and to consider the potato (see below on selection in early
quality trait of storage root flesh colour breeding stages). Most importantly, farmers
in the breeding process. Main emphasis in use more criteria and characters to select
breeding is given to: (1) material exchange sweet potato clones than do breeders in FPB
at the seed and clone level, (2) exchange of (Gibson et al., 2008). In such a situation,
information, knowledge and results from the risk of FPB is to ignore characters that
breeding trials by annual meetings, reports are important for good overall performance
and back-stop visits, and (3) a sweet potato of a clonal variety. However, in the study
breeding research and training build up of Gibson et al. (2008) in three provinces in
on the needs shaped among discussions Uganda, the most important characteristics
between NARS and CGIAR breeders. This for selection by farmers in early selection
has resulted in an additional aim to build stages were common to those used by
breeders, as were their relative weighting of a breeding programme (Gabriel and

characters, namely: (i) good root yield and Torrez, 2000; Manu-Aduening et al.,
big roots > (ii) SPVD tolerance or resistance 2006; Gibson et al., 2008);
> (iii) tolerance to drought, attractive root (ii) selection by farmers, mainly by
colour prior to cooking, straight root visual screening, is more efficient in
shape and orange- or yellow-flesh storage- earlier stages than in later stages of
root colour, and finally > (iv) tolerance to the breeding programme, which can
weevils. Characters of storage roots after also be explained by the larger genetic
cooking were not determined. It should be variation in early selection stages
noted that in later selection stages (FVS) the compared with later stages in breeding
priority list of farmers or relative character clonally propagated crops; and
weights changed, namely: (i) good root (iii) so far, selection by farmers at early
yield, big roots, drought tolerance, sweet selection stages has only been applied
and mealy roots after cooking > (ii) early to a sample of the genetic variation
root maturity, continuous root yield for generated by FPB in crossing pro-
piecemeal harvesting, and weevil tolerance grammes and it must be more efficient
> (iii) long root storage in the soil, extensive to expose the full genetic variation to
foliage, tolerance to caterpillars (Acraea farmer selection in the breeding pro-
acerata), marketability, attractive colour of cess.
storage roots prior to cooking, and non- In Sections 13.2 and 13.3 we suggest
fibrous roots after cooking > (iv) followed how this can be done in a cost-efficient way
by a group of characteristics with very low and without losing time in the breeding
weights, such as good root yield in poor process. However, doubts remain as to
soils, good vine establishment, tolerance whether farmers can efficiently use and treat
to rats and other vertebrates, non-sappy large amounts of true seeds and true seed
and no loss of taste in storage roots prior plants, which often appear in quite different
to cooking, soft texture, nice looking at the amounts per cross combination and have
table, nice flavour and easy or quick to cook quite different performance compared
storage roots. For some characters (mainly with plants grown from vegetative plant
biotic pressures, i.e. SPVD and weevil parts. It might be more useful that plant
tolerance; Gibson et al., 2008) there were breeders germinate seeds and multiply for
clearly different weights given to characters each family a reasonable numbers of clones
in different provinces, which might reflect so that farmers can select clones in small
local biotic challenge. Moreover, farmers plots comprising a few plants (2 to 4 per
used more attributes (51 attributes) than genotype). A further advantage of this is that
scientists and breeders (11 attributes) the breeder can use the frequency of selected
to describe and distinguish varieties. To clones per family by the farmer as additional
what extent this is important is not clear; information to identify appropriate parents
however, it might show the importance to for recombination. However, we think
farmers of distinguishing varieties. that the selection of parents in breeding
To summarize: clonally propagated crops should have a
(i) not surprisingly, farmers have the participatory component, but should be
ability to select successfully both in mainly carried out by the breeder due to
the early and later breeding stages of the genetics (see Section 13.2) and statistics
(see Section 13.5) involved in appropriate are eaten fresh, or are only boiled or
choices of parents in breeding clonally roasted, and when they are processed this
propagated crops. is often carried out at the household level.
To consider the range of needs, Exceptions are sugar cane, fruit crops used
preferences and adaptation requirements for the juice industry, and to certain extent
for the large number of clonally propagated root and tuber crops (potato, cassava and
crops is out of scope in this chapter. Here sweet potato) when they are used for the
we want to give the principles of breeding starch, alcohol or biofuel industries. In
clonally propagated crops and how PPB can resource-poor environments, yields and
be carried out or linked into these breeding yield stability with low input are a priority,
programmes. The breeding objectives in addition to consumer acceptability. As
and methods will be considered for four has been mentioned above, a major factor
agricultural crops in more detail at the that determines yields, yield stability and
end of this chapter, namely: potato, sweet adaptation are pests and diseases. The most
potato, cassava, and banana or plantain. important pests and diseases of important
Table 13.1 gives the plant parts used for clonally propagated crops in agriculture
propagation, the world production and by eco-geographical region are given in
the area harvested, as well as the polyploid Table 13.2, together with the most important
level of the most important clonally quality characteristics.
propagated crops in agriculture. Obviously, Most clonally propagated crops are
quality characteristics determined by polyploid (Table 13.1). An exception
consumer preferences and market needs is cassava, which can be considered as
are key characteristics for breeding clonally a polyploid behaving like a diploid (see
propagated crops, because many of these below). Polyploidy is an important aspect
TABLE 13.1
Data on the 11 most important clonally propagated crops on a global basis
Species Planting material World production† Area harvested† Polyploidy
Potato Sprout tubers 315 ×106 t 18.8 ×106 ha 2x, 3x, 4x, 5x
(Solanum tuberosum)
Cassava Hardwood cuttings 226 ×106 t 18.6 ×106 ha 2x
(Manihot esculenta)
Sweet potato Sprout cuttings 124 ×106 t 9 ×106 ha 6x
(Ipomoea batatas)
Yam Root tubers 51 ×106 t 4.6 ×106 ha 3x–10x
(Dioscorea spp.)
Taro Corms 12 ×106 t 1.8 ×106 ha 4x
(Colocasia esculenta)
Sugar cane Cane stalks 194 ×106 t ‡ 20.4 ×106 ha 8x
(Saccharum officinarum)
Banana and Plantain Corms 105 ×106 t 9.6 ×106 ha 3x
(Musa × paradisiaca)
Citrus fruit Bud stick grafting on 89 ×106 t 5.6 ×106 ha 2x, 3x+1, 4x-3
(Citrus spp.) rootstocks
Grapes Hardwood cuttings 69 ×106 t 7.4 ×106 ha 6x
(Vitis vinifera)
Apple Bud stick grafting on 64 ×106 t 4.8 ×106 ha 2x, 3x
(Malus pumila) rootstocks
Strawberry Adventitious shoots 4 ×106 t 0.26 ×106 ha 8x
(Fragaria grandiflora)
NOTES: † FAOStat 2006 at faostat.fao.org, ‡ Sucrose production.
TABLE 13.2
Quality characteristics, pests and diseases by production zone of the 11 most important clonally
propagated crops in agriculture and horticulture
Major production Quality characteristics Pest and diseases
zones
Potato (Solanum tuberosum)

Tropical highlands Various fresh consumption traits, high Late blight (Phytophthora infestans), cutworms (Agrotis
iron and zinc contents, adaptation to spp.), potato tuber moth (Phthorimaea spp.)
various micro-climates
Tropical lowlands More uniform fresh consumption traits, Bacterial wilt (Pseudomonas spp.), Early blight
high iron and zinc contents, extremely (Alternaria solani), Root-knot nematode (Meloidogyne
short crop duration (<80 days) spp.), viruses (Potato leaf roll virus (PLRV), Potato virus Y
(PVY), etc.), year round aphid pressure
Temperate zones Various fresh consumption traits, Late blight, cyst-forming nematodes, (Globodera spp.),
high starch for industrial use, various potato virus diseases (PLRV, PVY, PVX, etc.)
processing traits (chips, French fries)
Cassava (Manihot esculenta)
Humid tropics Cooking quality, elevated provitamin A Bacterial blight (Xanthomonas axonopodis) in Asia,
content for human consumption with Africa and the Americas, Frogskin disease (CFSD) in the
low HCN content, high DM for industrial Americas
uses
Drought-prone Cooking and processing (fried cassava, African cassava mosaic (CMD) virus and Cassava
tropics gari, fufu) quality, elevated provitamin brown streak disease (CBSD) in Africa, Green mite
A content for human consumption with (Mononychellus tanajoa) and mealybugs (Phenacoccus
low HCN content, high DM for industrial spp.) in Africa and the Americas
uses
Sweet potato (Ipomoea batatas)
Humid tropics High DM WFSP and OFSP Extreme Sweet potato virus disease (SPVD), especially in
eastern Africa
Drought-prone Elevated DM WFSP and OFSP, and clearly Sweet potato weevils (Cylas spp.) and SPVD to a lesser
tropics non-sweet in West Africa extent
Tropical highlands Elevated DM Alternaria spp. and SPVD to a lesser extent
Temperate zones OFSP with low DM and WFSP with high Root-knot nematode (Meloidogyne spp.) and SPVD to a
DM (both with medium sugar content) lesser extent
Yam (Dioscorea spp.)
Humid tropics Thirteen species with regional Yam tuber beetles (Heteroligus spp.) and Anthracnose
importance (main species D. rotundata), (Colletotrichum spp.), especially in West Africa, Yam
majority in wet hot tropics, but nematode (Scutellonema bradys), Root-knot nematode
D. abyssinica, D. alata and D. esculenta (Meloidogyne spp.) and Shoe string virus disease
also in dryer regions due to dormancy
of tubers; growing time and taste varies
extremely among species (some are
poisonous and must be cooked)
Taro (Colocasia esculenta)
Humid tropics Colocasia cultivar groups: (1) one large Corm and root rots (caused by Pythium spp.,
corm with few cormels; and (2) several Phytophthora spp., Rhizoctonia spp. and Erwinia spp.)
small cormels. Genotypes have very and Dasheen mosaic virus (DMV) across world regions,
different shelf lives (dormancy period) Taro blight (Phytophthora colocasiae) and Taro beetle
and some require excessive processing (Papuana spp.), especially in the South Pacific
before edible
Sugar cane (Saccharum officinarum)
All regions Weight of canes, sugar content, juice In the past, virus diseases were most important; today
purity, short or long vegetative times they play a subordinate role due to resistance breeding
and adaptation to photoperiod (non- and virus-free planting materials
flowering) Pineapple disease (Ceratocystis paradoxa), Red rot
(Colletotrichum falcatum), Smut (Ustilago citaminea),
Shoot and Internode Borer (Chilo spp.) in nearly all
regions
Humid tropics Yellow leaf spot (Cercospora spp.), Scale insect
(Melanaspis glomerata), Pyrilla (Pyrilla purpusilla)
Drought-prone Eye spot (Drechslera sacchari), Whitefly (Aleurolobus
tropics and barodensis)
subtropics
Tropical highlands Leaf scald (Xanthomonas albileneans), Wilt
(Cephalosporium sacchari)
Major production Quality characteristics Pest and diseases

zones
Banana and plantain (Musa x paradisiaca)

Humid tropics and Bananas have a lower DM and higher Banana wilt (Fusarium oxysporum) especially in the
subtropics sugar contents (very narrow genetic Americas, Yellow sigatoka (Mycosphaerella musicola),
variation in triploid gene pool – ca. Black sigatoka (M. fijiensis) especially in Asia, Moko
30 cvs.) compared with high DM disease (Pseudomonas solanacearum), Bunchy top virus,
and starchy plantains (larger genetic nematodes such as Radopholus similes, banana root
variation in triploid gene pool – ca. 125 borer (Cosmopolites sordidus)
cvs.). Plantains are important staples in
Central Africa and some parts of South
America.
Citrus fruit (Citrus spp.). Cultivated citrus may be derived from as few as four species:
Key Lime (C. aurantifolia), Pomelo (C. maxima), Citron (C. medica) and Mandarin (C. reticulata).
All other “species” are hybrids
All regions Very different tastes and fruit sizes Strong rootstock influence (C. jambhiri, C. reshni,
(oranges — ca. 1100 cvs.; mandarins, Poncirus trifoliata) in adaptation to cold and resistance
lemons, pomelo). Citrus trees hybridize to Phytophthora root rot and virus diseases such as
very readily and new hybrids easily Tristeza, Porosis and Exocortis
maintained by apomixis.
Subtropics Citrus canker (Xanthomonas citri), Foot rot
(Phytophthora spp.), Melanose (Diaporthe citri), Blue
and green mould (Penicillium spp.), Tristeza virus,
nematodes such as Tylenchulus semipenetrans, fruit fly
(Bactrocera spp.)
Drought-prone Foot rot (Phytophthora spp.), Gummosis (Phytophthora
tropics spp.), Citrus scab (Elsinoe fawcetti), Tristeza virus,
Porosis viruses, fruit fly (Bactrocera spp.), citrus psyllid
(Diaphorina citri), moth species such as Ophideres,
Sphingomorpha, etc.
Grapes (Vitis vinifera) North American species: V. aestivalis, V. labrusca, V. rotundifolia
Temperate zones The North American species are Bunch rot (Botrytis cinerea), Downy mildew
of interest for the summer rainfall (Peronospora sparsa), Powdery mildew (Erysiphe
regions in the tropics because of their necator), vine moths (Eupoecilia ambiguella, Lobesia
disease resistance and minimal chilling botrana), eriophyid mite (Calepitrimerus vitis)
requirement – especially in crosses with
V. vinifera (better taste, better texture
of berries)
Drought-prone Downy mildew, powdery mildew, Anthracnose
tropics and (Elsinoe ampelia), beetles such as Popillia japonica,
subtropics thrips (Scirtothrips dorsalis, Thrips hawaiiensis and
Rhipiphorothrips cruentatus), grape root borer (Vitacea
polistiformis), bugs such as Lygocoris inconspicuous,
Grape mealybug (Pseudococcus maritimus)
Apple (Malus pumila)
Temperate zones There are large differences in Fireblight (Erwina amylovora), Apple rust
vernalization need among cultivars and (Gymnosporangium spp.), Apple scab (Venturia
several can be grown very successful in inaequalis), Plum curculio (Conotrachelus nenuphar),
Mediterranean climates. Apple maggot (Rhagoletis pomonella), Codling moth
Some cultivars in higher places in the (Cydia pomonella)
equatorial region if the leaves are
removed before beginning of bud
dormancy (stripping off, or chemical
defoliation)
Subtropics Apple scab, Powdery mildew (Podosphaera leucotricha),
Crown rot (Phytophthora cactorum), Apple crown gall
(Agrobacterium tumefaciens), Bitter rot (Glomerella
cingulata), root rots (Phytophthora spp.), Woolly apple
aphid (Eriosoma lanigerum), Apple sawfly (Hoplocampa
testudinea), Green apple aphid (Aphis pomi)
Tropical Highlands Fireblight, Crown rot, Woolly apple aphid
Strawberry (Fragaria ×ananassa)
Subtropics and Ancient cross of F. virginiana (8x) from Grey mould (Botrytis cinerea), Powdery mildew
temperate zones eastern North America and F. chiloensis (Sphaerotheca macularis), Strawberry blossom weevil
(8x) from Chile (Anthonomus rubi), European tarnished plant bug
(Lygus rugulipennis)
Sources: Kranz, 1978; Rehm and Espig, 1984; Mandal, 2006.
in crop evolution (as we have already seen hexaploids (six sets; 6x) (Tate, Soltis and
in potato) and has important consequences Soltis, 2005) – and species with even higher
in breeding clonally propagated crops. It is polyploidy levels are known (Table 13.1).
important to note that all ‘breeding lines’ The haploid level (one set; 1x) does not
or varieties of clonally propagated crops are occur as a normal stage in the life cycle of
homogenous (clone lines and varieties are a crop. However, haploid plants occur by
genetically fixed and as homogenous as non- spontaneous mutations, wide crosses and
segregating breeding lines or hybrids from anther culture (e.g. diploids are developed
breeding self-fertilized or hybrid crops). from tetraploid potatoes by pollination with
The homogenous clones are exact genetic specific clones of S. phureja and haploids by
copies of their mother plants, if mutations anther culture). Haploids are occasionally
are ignored. This is more or less obvious in used in FPB of clonally propagated crops,
potato, cassava or sweet potato field plots, especially potatoes (Hermsen and Verdenius,
or in fruit and tree plantations, provided 1973; Wenzel and Foroughi-Wehr, 1984). In
no genotype mixtures are observed. What crop evolution different polyploidy levels
is not directly obvious to an observer originated from genome mutations and by
is that each clone line or variety in the hybridization between very closely related
field or plantation is a highly heterozygous species. Autopolyploids and allopolyploids
hybrid (clone lines and varieties are highly include wheat (Triticum durum and T.
heterozygous hybrids comparable with aestivum), canola (Brassica napus) and cotton
heterozygous hybrids developed in hybrid (Gossypium spp.), and nearly all clonally
breeding). It should be noted that due propagated species are autopolyploids. The
to polyploidy, clonally propagated crops homologous chromosomes in autopolyploids
are usually more heterozygous than those are similar enough that multivalents of the
diploid crops in which hybrid breeding same homologous chromosomes are formed.
is applied. The difference between “clone Doubling of chromosomes occurs if the
hybrids” and “seed hybrids” such as maize spindle poles are not developed when the
is that the first are propagated by asexual nucleus is dividing chromosomes in mitotic
reproduction and the latter are developed and meiotic cell division. There are several
by sexual reproduction. possible outcomes of abnormal meiosis.
Natural formation of 2n gametes was
13.3 POLYPLOIDY most important in evolution of cultivated
General knowledge about polyploidy is Solanum species, and the formation is mainly
required to get an understanding of breeding determined by one recessive gene (Watanabe
clonally propagated crops. Polyploidy has and Peloquin, 1988), so this character can
a strong effect on the performance of clones be used in breeding potato. Polysomic
as well as the parent–offspring correlations. inheritance is sensitive to disorders and
A polyploid genotype contains more than therefore autopolyploids often have reduced
two homologous sets of chromosomes in fertility, and occasionally they are completely
the nucleus of somatic cells. According to infertile and propagate only asexually.
the number of chromosome sets in the Multiple chromosome sets occur
nucleus we distinguish different polyploid spontaneously in nature from 2n gametes
types: triploid (three sets; 3x), tetraploid and can be induced artificially by colchicine
(four sets; 4x), pentaploids (five sets; 5x), (an alkaloid of autumn crocus, Colchicum
autumnale). In the case of diploid plants breeding behaviour of allopolyploids is

(2x), this leads to tetraploid plants (4x). An very similar to diploids. The formation
example is the evolution of S. tuberosum spp. of bivalents or multivalents appears to be
andigena (4x) from cultivated S. stenotomum genetically determined, e.g. without the
(2x) (Hawkes, 1979). Hybridization of gene (ph) on chromosome 5B in poly-
diploid and tetraploid plants forms triploid ploid wheat (Triticum durum and T. aes-
plants (3x) and by a further doubling of tivum), homologous chromosomes form
chromosomes hexaploid plants (6x) are multivalents. This gene is relatively new in
formed. An example is hexaploid I. batatas, the evolution of wheat (Dhaliwal, 1977).
which probably evolved by genome mutation Among clonally propagated crops, cas-
and hybridization, because the sweet sava (Manihot esculenta) is considered to
potato genome (6x) consists of two closely be a diploidized allotetraploid, which also
related sets of chromosomes (B1B1B2B2), was formed recently in the evolution of
of which one is duplicated (B1B1B2B2B2B2) plants (Nassar, 2000). Indications for this
(Shiotani and Kawase, 1989; Austin and are: (i) the high chromosome number (2x
Huamán, 1996). Many important clonally = 36) of all Manihot spp. (other Euphorbia
propagated crops are triploids (3x), such as have basic chromosome numbers within
the economically important genotypes of the range of six to eleven); (ii) natural
banana and plantain (Musa × paradisiaca). hybridization occurs among Manihot spe-
The triploid banana and plantain groups cies and crossing barriers appear to be
evolved in two different ways by genome weak; and (iii) M. esculenta shows meiotic
mutation and hybridization: in the case irregularities, such as terminal non-pairing,
of banana, from one diploid wild species multivalent associations and repetition of
M. acuminata (AA) to form the triploid chromosome types, which results in low
banana group (AAA), and in the case of fertility of parental combinations.
plantain, from two diploid wild species: Polyploid plants usually have larger
M. acuminata (AA) and A. balbisiana (BB), plant cells, larger and stronger plant
forming the triploid plantain group (AAB) organs, greater height and increased
(Simmonds, 1962). Many FVs in the banana biomass production. In nature, polyploid
and plantain group evolved only by somatic plants tend to succeed in new habitats.
mutation, because triploid banana and In breeding, the tallest and best thriving
plantain are infertile. However, breeding plants are selected, so that, unintentionally,
triploids is possible by working with two many crops have been bred to a higher
gene pools, one which is diploid and the level of ploidy. However, as chromosome
other which is tetraploid, such as using gene number increases, the increase in biomass
pools of M. acuminata and M. balbisiana on production becomes successively less,
a diploid and tetraploid polyploidy level to and production decreases above a specific
develop new triploid banana and plantain optimum biomass. This optimum differs
varieties. from species to species. In autopolyploids,
In contrast to autopolyploids, the this advantage of increased vigour is
genome in allopolyploids differs so much associated with the disadvantage of increased
between hybridized species that only biva- meiotic disorders during the formation of
lents of homologous chromosomes of multivalents. This is the reason why the
the parental genomes can be formed. The harvest in many important autopolyploid
crops is represented by vegetative plant gametes), has been proposed for breeding
parts. Most polyploids display heterosis tetraploid potatoes (Ortiz, 1998). Polyploidy,
relative to their parental species, as well as heterozygosity and heterosis make the
relative to inter-gene-pool crossings within selection of good parents in population
a species. A polyploid population contains improvement of clonally propagated crops
three, four, five, six or more alleles at each very difficult. A good parent generates
locus. Hence, considerably more effects large genetic variation around a high family
due to dominance and epistasis are possible, mean. Cross-prediction and inter-gene-pool
and the genetic variation due to dominance crosses are very important in population
and epistatic effects in polyploidy crops improvement of clonally propagated crops.
is very large compared with the genetic This aspect of clonal breeding is often
variation caused by dominance and epistatic neglected and this might be the reason for
effects in diploid crops. For this reason, the low level of breeding progress in many
the performance of clonally propagated clonally propagated crops. In contrast
crops is mainly determined by heterosis. to population improvement (selection of
Usually in breeding of clonally propagated superior parents – see Section 13.7 below),
crops, an F1 clone hybrid is crossed selection within a given genetic variation
with another F1 clone hybrid, so that for variety development is relatively easy in
the offspring shows extremely extensive clonally propagated crops (discard inferior
segregation. In parent–offspring studies it material). All the genetic advantages of
is possible to determine mid-parent and clonally propagated crops can be used for
mid-offspring heterosis, as well as the best- variety development, and the genotype
parent mid-offspring heterosis (similar to finally released is in the hands of the breeder
the assessment of heterosis in a hybrid immediately after the initial crossings.
breeding programme of diploid crops—see A clonally propagated crop that has no,
Chapter 11). In polyploids, more than one or nearly no, sexual reproduction is close
allele per locus is transferred in gametes to a dead end in evolution and breeding.
to the next generation, so that, in contrast Genetic variability can only accumulate
to diploids, the genetic variation due to by mutations. However, this source of
dominance determines the response to new variation has often been used to find
selection in population improvement as enhanced types of fruits and ornamentals
long as the population is not in equilibrium (van Harten and Broertjes, 1988).
(after recombining parental material in Nevertheless, the main source of generating
controlled crossings, a population is usually new variation in clonally propagated crops
not in equilibrium). In tetraploid potato is sexual reproduction. Owing to a more
populations that are not in equilibrium, or less regular meiosis in polyploids with
one-third of the dominance variance is an even number of chromosome sets
exploitable for selection progress when (4x or 6x), sexual seed production and
selection takes place on the female and male generation of new genotypes is possible.
sides (Wricke and Weber, 1986; Gallais, Nearly all clonally propagated crops, e.g.
2004). The exploitation of the dominance potato, sweet potato and cassava, are cross-
variance in population improvement, in fertilized crops in combination with self-
combination with the selection for different incompatibility. Incompatibility alleles
levels of ploidy (using the inheritance of 2n are the reason why specifically sought
after cross combinations are difficult to and doubled-triploid production. The

realize, and seeds from controlled crossings populations developed from seeds are
can have a very high value in clonally again formed by very different and highly
propagated crops. heterozygous genotypes, which do not
exchange genetic material. Each seed plant
13.4 GENERAL BREEDING SCHEMES grown in the so-called seedling nursery can
The general principle of breeding clonally be considered a potentially new variety.
propagated crops is to break normal clonal This is the basis for selection. The selection
propagation by introducing a crossing step, between clones is described most often
which culminates in sexual seed production in plant breeding textbooks as a process
and genetic variation. After the genetic conducted in several steps (Figure 13.1).
recombination, all subsequent propagation The breeding scheme illustrated in
steps are asexual in nature and done by Figure 13.1 is straightforward and it is
clonal propagation. Nearly all clonally most often interpreted as requiring clonally
propagated crops are polyploid and cross- propagated crops to be bred sequentially
fertilized species. A more or less regular in several steps over several years. The dia-
meiosis is possible in polyploids, if the gram implies that there are two parents
number of chromosome sets is even, as being crossed, followed by five subsequent
in tetraploids (4x) and hexaploids (6x). selection steps in time (one selection step in
The parents in cross combinations are seed plants and four selection steps in clone
highly heterozygous hybrids, with the plants). This is misleading. First the breeder
exceptions of inbreeding lines generated must work with many parents (further
by self-fertilizations or doubled-haploid details about number and size of crosses
are given in Section 13.5). Second, there is
no further genetic development in clonally
FIGURE 13.1
The general breeding scheme
propagated crops as one moves between
for clonally propagated crops selection steps. The selected D-clone in
Figure 13.1 is genetically identical to the
Crossings true seed plant the selected D-clone derives
from. Provided that the true seed plant can
True seed plants be cloned in large quantities, it is theoreti-
cally possible to test the population with
A-Clones adequate accuracy in the first year to select
the ‘best’ genotype.
B-Clones Selection among true seed plants is made
for tolerance and resistance to pathogens.
C-Clones However, often no selection between plants
grown from seed is made by the breeder.
D-Clones Nevertheless, natural selection occurs during
germination, and should not be completely
Propagation avoided, because genotypes difficult to
germinate delay the breeding programme.
The main reasons for no selection in the
Source: modified from Becker, 1993. seedling nursery are: (i) plants grown from
seed often differ considerably from plants years. It is obvious that the wide range of
raised from vegetative planting material, quality preferences and the numerous pests
(ii) the plants raised from seeds are normally and diseases in each clonally propagated
grown in pots in greenhouses, and for crop and their interaction with genotypes
most traits this is not representative of justifies decentralization and participatory
field conditions, and (iii) a single plant approaches. However, the better simulation
evaluation is usually not appropriate, with of the final target environment realized
the exceptions of susceptibility to highly with FPB justifies a stronger PPB approach.
aggressive pathogens. In field crops, an Many advocate PPB because the stress and
important factor is interplant competition. marginal field conditions of resource-poor
A genotype must be tested under conditions farmers are not adequately simulated by
that simulate the field conditions in practice. FPB (see also below). In this context the
For this reason several plants of each two clear advantages of breeding clonally
clone are tested in plots in blocks under propagated crops should be pointed out:
homogenous field conditions. The aim is an (i) no genetic changes occur in genotypes
unbiased comparison of genotypes within after seed has been produced; and (ii) the
blocks. The number of plants per plot and total genetic variation of genotypes
the plot size depends on the crop as well as (comprising the genetic variances due to
the breeding stage. Fruit trees and perennial additive, dominance and epistatic effects)
shrubs are tested in larger plots with fewer can be exploited by selection. For these
plants than potatoes, and these again are reasons, only the genotype×environment
tested in larger plots than cut flowers. Early (G×E) interaction and the plot error must be
selection stages (A-clones and B-clones) are considered (and reduced by testing in several
tested in smaller plots than later selection environments) to identify the best clone.
stages (C-clones and D-clones). The amount
of planting material at each breeding stage is 13.4.1 Early breeding stages and PPB
determined by the propagation coefficient In the general breeding scheme
of the crop. For example potato has, (Figure 13.1) each surviving seed plant
among clonally propagated crops, a very is cloned to be raised as A-clones in
low propagation coefficient of about ten, observation plots (visual screening of
whereas sweet potato has a relatively high general clone performance), or evaluation
propagation coefficient of between 30 and plots (recording of data on specific traits
90 (depending on the field propagation of each clone). Figure 13.2 shows the
method used). This is one factor why potato planting of sweet potato A-clones. The
breeding is relatively slow (about eight to plot size of A-clones is usually a single-row
ten years from cross to variety release). plot comprising 3 to 5 plants. The trial is
Breeders do not breed for a single conducted with no replications. It is open
environment; they breed for a range to discussion whether A-clones should
of environments. Hence, the field be evaluated at two locations. Selection
evaluations must simulate the range of theory results show that it is nearly always
target environments. For this reason, and the best resource allocation to test as many
depending on the propagation coefficient, clones as possible at one location, without
the clones are tested in plots, in homogenous replications (Wricke and Weber, 1986).
blocks, at several locations and for several Many breeders use only one location at the
FIGURE 13.2
Planting early selection stages of sweet potato for the
accelerated breeding scheme in San Ramon (one of four locations)
early breeding stages due to the restrictions a marginal or hot-spot environment (for
of the propagation coefficient and breeding drought, salinity, biotic challenge, etc.) can
budget. However, there are several reasons be discarded, or at least considered with
to test A-clones at two locations: (i) a trial caution in good environments.
at one location can be lost (e.g. extreme A-clones are only selected for highly
weather conditions) and then a full breeding heritable traits such as general performance
step and population is lost; (ii) trials at only (growth type; tuber, root or fruit size, shape
one location are of little value (the G×E and colour), resistance to pests and diseases,
interaction cannot be separated from the harvest index, dry matter and nutritional
genotypic effect); and (iii) the response to quality. Breeders nearly always conduct
selection is still very close to the optimum a visual selection at the A-clone breeding
in a wide range of scenarios, including stage. However, it can be questioned if the
the scenarios where A-clones are tested A-clones selected by the breeder match
at two locations (Grüneberg et al., 2004). farmer needs and would be selected by
Moreover, information from contrasting farmers. With two locations, one location
environments can be combined if the can be easily evaluated by farmers in a
breeder tests A-clones at two locations. PPB approach, while the other location
For example, clones that clearly fail in is used by the breeder. It should be noted
that visual selection of general performance nutritional quality (starch, vitamins and
can also be an efficient indirect selection micronutrients by fast through-put analysis
for yield. In sweet potato, we observed methods) (Hartmann and Buning-Pfaue,
among several thousand A-clones grown in 1998; Lu, Huang and Zhang, 2006; Zum
1 m-row plots at three locations a heritability Felde et al., 2007; Bonierbale et al., 2009).
for yield of about h2 ≈ 0.4 (harvesting and In the next season, B-clones—also
recording all A-clones for yield at all three called “promising clones”—are planted in
locations). It was considered as ‘useless larger plots in 2 to 3 rows with planting
work’, because a visual selection at the material obtained from selected A-clones.
first location resulted in a nearly common The B-clone trials are still conducted
set of selected clones and a heritability without replications, but generally at two
for yield of about h2 ≈ 0 in the selected or more locations. The B-clone stage is
fraction. This was demonstrated for two usually the beginning of selection for low
different breeding populations grown in heritability traits such as yield, biomass
two different seasons, so that the breeding and yield stability. The determination of
scheme was changed. Only those clones stability parameters such as the slope of
that have passed the visual selection step at the regression line and deviations from
location 1 are harvested and considered for regression (Fox, Crossa and Romagosa,
storage root quality evaluations at location 2 1997) requires at least three locations.
and 3. However, relying on visual selection However, it should be noted that stability
in early breeding stages requires a person parameters from less than 6 environments
who is very experienced with sweet potato. are still of little value. As mentioned above,
We think farmer participation at the visual a strong justification for PBB is that stress
selection stage in early breeding stages and marginal field conditions of resource-
is essential to avoid genotypes entering poor farmers are not adequately simulated
later breeding stages with characteristics by FPB (Ceccarelli, 1994). Cross-over G×E
(storage root size, shape, form, colour, etc.) interactions occur, and what appears to be
unacceptable to farmers. As described above, good in resource-rich environments often
farmer preferences vary substantially both does not perform well in resource-poor
within and between regions, and the visual environments. This has also been clearly
selection can be conducted by independent observed in sweet potato, and outstanding
farmer groups. The advantages of PPB are clones for resource-poor environments were
very obvious in the early selection stages discarded by FPB (e.g. the clone SR92.499-
of clonal breeding, in which large numbers 23; Grüneberg et al., 2005). Usually, but not
of fixed genotypes must be screened for always, the response to selection in poor
many highly heritable traits. PPB in the production environments is smaller than in
early selection stages has been successfully good production environments. The genetic
applied in potato (Gabriel and Torrez, variance is smaller while interaction and
2000), cassava (Manu-Aduening et al., error are larger, so that the performance of
2006) and sweet potato (Gibson et al., individual clones becomes more difficult
2008), and by working with two or three to distinguish. However, outstanding
locations it can be linked into FPB, in genotypes with different growth types
which selection is conducted for traits that adapted to resource-poor environments
cannot be evaluated by farmers, such as cannot display their full potential if FPB
does not test in such environments. Taking 3- to 5-row plots. All important agronomic
sweet potato breeding as an example again, traits are determined, including taste and
yield stability is associated with harvest post-harvest characteristics. Furthermore,
index (Grüneberg et al., 2005). Under it merits determination of the above-
drought stress, good performing sweet mentioned stability parameters: (i) slope
potato clones have a harvest index of about of the regression line, and (ii) deviations
0.5 (on the basis of fresh matter storage from regression, as well as conducting an
root yields and total fresh matter biomass Additive Main Effect and Multiplicative
yields). Vine production is of considerable Interaction (AMMI) Analysis in those
importance to farmers to obtain sufficient cases where the regression model does
planting material for the next growing not fit (Fox, Crossa and Romagosa, 1997).
season. Clones performing well in resource- Usually, a clone is considered to have stable
rich environments usually fail in drought- performance if the slope of the regression
stress environments due to insufficient vine line is close to 1, and the deviations from
production rather than to unacceptable the regression line are small. An important
storage root production. At the same question in later breeding stages is that
time, outstanding clones in drought-stress of how many locations and how many
environments show a strong increase in vine replications to use. With more locations and
production with medium storage root yields more replications, the estimation of the yield
when grown in environments with good performance of clones is more reliable. At
water supply (Andrade, unpublished). The the same time, for a given testing capacity,
selection of genotypes with desired growth increasing the number of locations and
types or desired sink–source allocations replications results in fewer clones being
in marginal environments requires that tested. Generally, the gain from increasing
breeders evaluate the breeding population the number of replications is less than
in such an environment; this characteristic that obtained by increasing the number of
cannot be determined in a resource-rich genotypes and locations. Investigations of
environment. Here we suggest linking this problem in selection theory have led
the evaluation in a marginal environment to a recommendation to conduct advanced
with the visual selection in early breeding clone trials still with no replications but in
stages. All clones that fail in the marginal the maximum number of environments that
environment (i.e. extreme reduced storage can be managed by the breeder (Utz, 1969,
root production or vine production) are cited in Wricke and Weber, 1986). However,
eliminated from all other selection steps. many scientists are still very reluctant to
conduct trials without replications. Since
13.4.2 Later breeding stages and PPB the fixed costs of experimental stations are
At the beginning of the C-clone and high, it is usually an advantage to (i) create
D-clone selection stages the breeding ‘artificial environments’ on experimental
population has been reduced to between stations (by running part of a station without
30 and 300 clones. While the number of fertilizer or with less irrigation) and (ii) to
clones in later selection stages is further go on-farm to evaluate clones with farmers,
reduced, those selected clones are tested i.e. PVS. However, nearly all of the initial
in more environments and in replications. genetic variation in the breeding population
The plots for C-clones and D-clones are has been discarded at later breeding stages,
so that specific characteristics needed by number of genotypes at the first stage, to

farmers and consumers are often no longer the maximum of the available breeding
present in advanced or elite clones if they capacity; (ii) to use a high selection intensity;
had not been considered at earlier breeding and (iii) to use as many environments as can
stages. be managed at each breeding stage (Wricke
and Weber, 1986). Replications are of minor
13.5 MODIFICATIONS OF THE GENERAL importance and should only be used at the
BREEDING SCHEME final breeding stages. These characteristics
The general principle for breeding clonally of the optimum in multistage selection
propagated crops presented above is very for clonally propagated crops led to the
simplified. In practice, it is more or less suggestion of using an accelerated breeding
modified. The differences can be large, scheme (ABS) for clonally propagated
depending on the crop, country and breeder. crops in sweet potato breeding (Grüneberg,
For example, resistance or tolerance can unpublished).
already be determined at the true-seed ABS responds to the frustration that
plant stage by eliminating infected plants it takes on average seven or eight years
from the seed nursery. Potato breeders from a cross until variety release. Donors
usually try to obtain only a single tuber are also reluctant to invest in breeding
from each true seed plant to start selection when concrete outputs take so long to
with single plant tests. Clone selection in materialize. ABS uses the simple fact that
shrubs and fruit trees uses fewer plants in breeding clonally propagated crops each
per row and fewer selection stages. Potato true seed plant is already a potential variety,
breeding uses more selection stages due to with the advantages of sweet potato having
the low propagation coefficient of potato. a very short crop duration (3 to 4 months)
However, there is a common question in and a high propagation coefficient (up to 90
all the different breeding schemes: How cuttings per plant within 3 to 4 months). ABS
many genotypes should be selected at each overturns the general principal breeding
selection stage? In selection of breeding scheme of clonally propagated crops by:
clonally propagated crops this can be easily (i) crossing and multiplication; (ii) early
determined using selection theory. There selection stages; and (iii) late selection
is an optimum number of clones, locations stages. Everything that can be implemented
and replications at each selection step for simultaneously in these three stages and
a given test capacity. Fortunately, the area years is done simultaneously in different
around the optimum is flat and deviations environments. However, to reduce labour,
from the optimum do not have large effects, every clone that has not met a desired target
as long as the deviations are not strong. for a character in the first environment is
To select between 5 and 20 percent of the discarded and not considered (harvested)
total number of clones at each step is still in the second environment, and the same
close to the optimum. However, in the for characters evaluated in the second
wide range of practical breeding situations, environment, and so forth. In selection
the optimum has always been found in the theory, this multi-trait selection procedure
direction of higher selection intensities, is designated ‘independent culling’ and it
more so than most breeders intuitively is the procedure also used to optimize
realize. It is important: (i) to increase the multistage selection procedures (Cochran,
1951; Wricke and Weber, 1986). In ABS, from two hybrids cannot be reproduced
independent culling is conducted: (i) in a by crossing the hybrids again) without a
poor resource environment where clones system that maintains and provides at least
undergo visual selection; (ii) only those some healthy planting material.
clones passing the first selection step are Numerous viruses, bacteria and fungi
harvested in environments 2 and 3 to are transmitted by vegetative planting
determine yield and quality of selected material. Viruses are particularly important,
good performance clones over all traits and because viral diseases cannot be controlled
environments (index values are determined chemically. Viruses are spread by vectors,
by the Pesek-Baker index (Pesek and Baker, most often aphids and whiteflies. The
1969) to assist the breeder in their selection traditional maintenance of varieties and
decisions); and (iii) only those clones that production of healthy planting material
have passed the second selection step are includes protecting the base plants of
harvested in environment 4, where clones varieties in greenhouses or under nets, and
were already planted in season 2 in a farmer’s to prevent the development of a vector
field under high SPVD pressure in a third population by intensive use of insecticides.
selection step to select for SPVD tolerance. The base material is also termed ‘mother
About 300 sweet potato clones enter the later plants’. However, under these conditions,
breeding stages. In two subsequent seasons only 20 to 200 plants of each variety can
and two selections steps, 4 to 5 clones are be maintained, and planting material must
finally selected for variety release (first be produced in the field. These clones in
season: 300 clones, three environments, two the field for producing healthy planting
plot replications and 5-row plots; second material are the so-called S-clones, because
season: 40 clones, 16 environments, two planting material is usually called seed in
plot replications and 5-row plots). This is clonally propagated crops. Healthy S-clone
carried out in cooperation with NARS and production is supported by (i) application
farmer groups. of insecticides against vector populations
(monitoring by yellow cards); (ii) choosing
13.6 MAINTAINING VARIETIES AND locations for S-clone production that
S-CLONE MULTIPLICATION are out of range of vector populations
As a result of clonal propagation, (i.e. locations close to the sea or in cool
maintaining varieties should not be highlands); and (iii) removing all visibly
difficult. Genetic changes in varieties do infected plants from S-clone fields.
not occur by undesired crossings nor The detection of virus infections has
by segregation, and mutations are rare. been simplified by use of the enzyme-linked
However, the opposite is the truth, and immunosorbent assay (ELISA) procedure.
maintaining clonally propagated varieties is The principle is a reaction between the
a difficult and expensive part of the breeding viruses in plants and antibodies against these
operation. The main reason is that in clonal viruses. The reaction is made visible by an
propagation through vegetative plant parts, enzymatic colour formation. In practice,
many more diseases can be transmitted some leaf sap is pressed out and the colour
compared with seed propagation. A new reaction is assessed on special test plates
variety will have no impact in practice, and coated with antibodies. In the case of sweet
even can be lost (a clone hybrid developed potato, the plants tested negative for viruses
are further grafted on an indicator plant certain discipline in S-clone production

such as Ipomoea setosa to confirm the on farm, the yield level remains low,
absence of viruses for sweet potato viruses. although virus-tolerant varieties with
In this way all maintained mother plants of good overall performance are available.
a variety are routinely screened, and only The most important factors for S-clone
virus-free mother plants are used for further production on farm are: (i) separating
propagation steps. Recently, techniques have S-clone production from cultivation
been developed to detect viral DNA and for production; (ii) removing all visibly
RNA directly by real-time polymerase chain infected plants in S-clone field areas; and
reaction (PCR) (Mumford et al., 2006). (iii) obtaining new, healthy planting material
However, the best option for maintaining at least occasionally from private or public
clone genotypes is to start from absolutely sources. Nevertheless, the private and
virus-free material. This is obtained by in public seed sectors are an important factor
vitro propagation of plants under sterile in production of clonally-propagated crops,
conditions, and these in vitro plantlets are but this topic belongs to integrated crop
the starting point for greenhouse and field and pest management (Salazar, 1996). The
propagation. In vitro plantlets are replacing breeder’s role in this context is to maintain
mother plants in the greenhouse, often by and provide virus-free starter material for
eliminating all greenhouse plants. If no the private and public seed sectors.
virus-free material is available, new virus-
free plantlets can be obtained by thermo- 13.7 SELECTION OF PARENTS AND
therapy and meristem culture. Meristems PREDICTION OF CROSS OUTCOMES
of very-fast-growing infected plants are The choice of parents is perhaps the most
virus free following proper heat treatment, important step in a breeding programme.
because viruses only start to enter older Many breeders make several hundred crosses
plant cells. However, this process requires each year and it is often observed that
considerable time (at least 18 months for in later steps of the breeding programme
sweet potato, and depends on the virus titre the best clones derive from one or only a
of the infected source plants). In breeding, very few crosses. Hence, there is a desire
virus-free material can be achieved by ger- to predict which cross combinations are
minating true seed in vitro and maintaining most promising. If this were possible, the
these true-seed plantlets in vitro until the efficiency of a breeding programme could be
final selection decision has been made. increased by reducing the number of cross
Distribution channels for clonally combinations and increasing the number of
propagated crops are well developed in genotypes from good cross combinations
temperate regions of the world. However, (produce more genotypes from within the
they are almost non-existent in most best families). In the situation where not
tropical and subtropical countries, although much is known about the performance of a
the pest and disease pressure is considerable cross, the number of combinations should be
higher than in temperate regions. S-clone increased to the maximum of the breeder’s
production in resource-poor environments capacity and the number of genotypes per
is nearly all in the hands of farmers, and cross should be kept small. The rationale
the health status of planting material is a underlying this is based on selection theory,
key factor in high farm yields. Without a which shows that if “the breeder has no
prior knowledge on the cross … the breeder are rarely used in practice. One suggestion
has to make as many crosses as possible”, is to determine the value of a parent on the
which is also minimizing the risk of raising basis of the offspring performance from test
genotypes with poor performance (Wricke crosses. Another suggestion is to work on
and Weber, 1986). As mentioned above, most a reduced polyploidy level, which has been
clonally propagated crops are polyploid and especially proposed for breeding tetraploid
highly heterozygous, so that dominance potatoes (Ross, 1986). However, the latter
and epistatic effects contribute considerably has been little applied in practice for
to clone performance. For this reason, it parental selection, but has often been used
should be assumed that not much is known to incorporate germplasm of wild Solanum
about the value of a cross combination species into advanced breeding populations
until it has been made and tested. This is in (Tarn et al., 1992). Parental selection on the
agreement with our observations in sweet basis of test crosses are made on a large scale
potato, where the correlation between in potato breeding programmes for long-
mid-parent and mid-offspring yields is low day, temperate climates (150 to 500 cross
(r ≈ 0.5). We currently recommend raising combinations per breeding programme,
10 to 20 genotypes per cross combination, cited by Ross, 1985). It has been observed
while increasing the number of cross that specific combining ability is nearly
combinations to the maximum possible as large as general combining ability, and
with the resources available. However, after in some cases specific combining ability
clones of these crosses have been evaluated, has been observed that is clearly larger
the good crosses should be repeated on a than general combining ability (Sanford,
large scale. An optimum for the number and 1960; Mullin and Lauer, 1966; Tai, 1976;
size of crosses can determined if estimations Killick, 1977; Veilleux and Lauer, 1981;
are available for the genotypic variance Gaur, Gopal and Rana, 1983, cited by Tarn
between crosses and within crosses, and the et al., 1992; Gopal, 1998; Kumar, 2004; De
non-genetic variance components (Wricke Galarreta et al., 2006). This is not surprising
and Weber, 1986). Breeders often generate a as long as potato breeders do not work with
large number of seeds in polycross nurseries, two clearly separate gene pools for variety
but in these only the female parent is development. In potato breeding for tropical
controlled. The correlation between parent and subtropical regions, heterosis and
and mid-offspring in breeding populations high general combining ability have been
derived from polycross nurseries is half observed between andigena and tuberosus
of mid-parent–mid-offspring correlation in gene pools in tuber-propagated potatoes
controlled crosses. and in true-seed potatoes (Enrique Chujoy,
Often the parents are chosen due to their pers. comm.). However, as long as these
performance per se. For theoretical reasons, gene pools are not improved on the basis of
this cannot be very secure in clonally- general combining ability separately from
propagated crops. Clone varieties are the complementary gene pool, such effects
highly heterozygous hybrids and usually cannot be exploited in the long term.
polyploids, so that segregation in crossings In sweet potato experiments we observed
is almost unpredictable. Therefore, for a long a mid-parent–mid-offspring heterosis of
time now, suggestions have been made for 84 percent among 48 cross combinations (or
better assessment of parents; however, they 184 percent if the mid-parent value is set to
100 percent). This is a clear indication that colour, as well as pest and disease resistanc-
the design of breeding schemes using the es), while others are selected simultaneous-
combining ability of two gene pools merits ly by aggregating characters into an index
investigation. Two breeding gene pools are (often an intuitively formed index, such as
available for sweet potato to test heterosis: score values for overall performance).
the Jewel Gene pool, developed mainly A parent appears to have a good overall
from North American varieties, and the trait performance if no trait is below the
Zapallo-SPK Gene pool, developed mainly population average. However, only in those
from South American and African FVs. cases where trait associations are close to
The value of a parent is nearly always zero or positive can it be expected that
determined by several characteristics. In parents with good performance over all
general, parents should be recombined with traits produce offspring in which each char-
a good combining ability and good per- acter has been improved. In parental selec-
formance over all traits. The PPB study in tions, negative trait associations can be very
Uganda (Gibson et al., 2008) underlines critical. Table 13.3 gives an example for
how many characteristics are important for sweet potato, in which DM shows a strong
good performance over all traits. Moreover, negative trait association with pro-vitamin
FPB also has the aim of improving nutri- A, iron and zinc concentrations, as well as
tional quality, especially pro-vitamin A, a moderate negative trait association with
iron and zinc concentrations (Pfeiffer and storage root yield. The associations in the
McClafferty, 2006) in potato, sweet potato, example are strong enough that under vari-
cassava, plantain and other crops. With an ous scenarios of multi-trait selection the
increasing number of characters, breeders breeding population is improved for yield,
operate with larger breeding populations, pro-vitamin A, iron and zinc, whereas the
as in potato and sweet potato. Aiming at DM of the population decreases.
only 30 genotypes finally selected, and In other words, the DM is changing
assuming 10 characters each, selected in in the wrong direction even though it
sequential selection steps with a selected was selected for improvement. These sur-
fraction of ten percent (1 out of 10), then prising undesired effects in the case of
300 000 000 000 genotypes would be needed sweet potato and DM improvement in
in the original base population. Populations connection with pro-vitamin A, iron and
of this size cannot be established in prac- zinc improvement was also observed for
tice. Moreover, even if the population size the Williams selection procedure and this
is extremely large, some desired combi- index selection procedure (Williams, 1962)
nations probably do not exist, such as comes very close to intuitive selection pro-
sweet potato genotypes with high yield, cedures used by breeders in which a weight
high SPVD tolerance, high DM and high is assigned to each trait on the basis of its
pro-vitamin A, iron and zinc concentra- economic importance. The only selection
tions). Often, breeding can only approach procedure that can monitor the response
the desired genotype in several steps of to selection in each trait is the Pesek Baker
recombination and selection. In practice, index (Pesek and Baker, 1969). However,
some characters are selected sequentially this index requires estimations of genetic
(especially where there is clearly a low- variance and co-variances, but the proce-
est acceptable value (tuber size, shape and dure ensures that parents are selected that
TABLE 13.3
Estimations of genetic correlations for yield, dry matter, total carotenoids, iron and zinc in sweet
potato storage roots of 24 megaclones and 26 advanced breeding clones grown in at two locations
in two replications
Storage root yield Dry matter Total carotenoids Iron
Megaclones (orange and white fleshed)

Dry matter -0.49
Total carotenoids -0.06 -0.54
Iron -0.26 -0.23 0.94
Zinc -0.22 -0.39 0.93 0.74
Advanced breeding clones (only orange fleshed)

Dry matter -0.54
Total carotenoids 0.55 -0.71
Iron -0.24 -0.07 0.14
Zinc 0.39 -0.20 0.37 0.53
develop populations in which traits are combining ability and improving gene pools
improved according to a ratio of desired on the basis of general combining ability
genetic improvements (so-called desired values (called reciprocal recurrent selection
genetic gains) given by the breeder. in maize breeding) are the most difficult
An alternative is the Elston index tasks in breeding; however, they can greatly
(Elston, 1963), in which the breeder can increase yield gains. At the same time, we
raise the threshold for the trait at risk by think that the visual selection of potential
modifying the lowest acceptable value for parents in a PPB approach should be used
each. This index can be easily applied in as additional information by the breeder. It
each replication and environment, so that should be noted that the work plan for both
index mean values for each genotype can the selection of parents for the next cycle of
be calculated together with other statistical selection and the early selection stages for
parameters (Grüneberg et al., 2005). variety development are always to a certain
We are aware of only one case in which extent in common. In sweet potato breeding
PPB has been applied for the selection of at CIP we use a combination of sequential
parents in clonally propagated crops. In and simultaneous index selection in early
the Cochabamba region of Bolivia, farmers selection stages (see also above): (i) visual
selected potato parents in an andigena selection by eliminating all genotypes that
population, which had been improved do not meet the lowest acceptable values
for agronomic performance and Late for each trait (this lends itself to PPB);
blight tolerance. Selected clones in this (ii) in the remaining selected fraction (about
population were used as parents with the 2 500 clones), apply index selection for
regionally grown FV ‘Waycha’ (Gabriel yield and nutritional quality traits using the
and Torrez, 2000) and the PPB approach Pesek-Baker index, with the square roots
included hand-crossing by farmers. We of variance components as desired genetic
think that the ability of farmers in the gains; and (iii) selecting for pest and disease
selection of parents is limited beyond a tolerance (mainly SPVD) in the remaining
selection of clone performance per se. Test selected fraction (about 300 clones) by
crosses, general combining ability, specific visual selection (this lends itself to PPB) and
ELISA. The remaining 100 to 200 clones 13.8 APOMIXIS

enter later breeding stages, but are also As mentioned earlier, the principle advantage
used as parental material for the next cycle of breeding clonally propagated crops is that
of recombination and selection. In such a each clone variety is fixed and maintainable.
breeding system, with one population, two However, this is associated with the
PPB steps can easily be applied. However, disadvantage of vegetative propagation.
a PPB approach is feasible also in inter- Diseases are easily transmitted and the
pool crosses linked with general combining maintenance of varieties and the production
ability improvement. Farmers select in of healthy planting material are expensive.
families (derived from recombining the The ideal propagation system for clone
two gene pools) in early generations for varieties would be vegetative propagation
variety development (as described above). by seeds. This ideal propagation system
The interesting information for the breeder exists in nature, and is called apomixis
provided by farmers could be the numbers (Nogler, 1984). Apomixis is the formation
of selected clones per family. With this of seeds without meiosis, and two forms are
information the breeder can focus only on distinguished: (i) agamogenesis (also called
those parents in the improvement of the gametophytic apomixis), in which the asexual
separate gene pools, which for the farmer embryo is formed from an unfertilized egg;
results in interesting cross combinations and (ii) adventitious embryony, in which
with the other gene pool. On top of this, the the asexual embryo is formed from nucellus
breeder can use the opportunity to apply tissue. Apomictically produced seeds are
the general combining ability concept. This genetically identical with the parent plant.
would be a very elegant PPB approach for The breeding work on apomictic species
selection of parents and cross prediction. is very difficult and requires developing
Although Hull (1945), in his fundamental population improvement by sexual
paper on reciprocal recurrent selection, reproduction and subsequent variety
proposed this for breeding clonally development by apomixis. Apart from
propagated crops, this method of clonal some forage (Miles, 2007) and citrus species
breeding is rarely found in practice. (Soost and Roose, 1996), apomixis is not
The topic has been considered in breeding used in plant breeding.
clonally propagated trees (e.g. Baudouin et The difficulty in breeding apomictic
al., 1997; Kopp et al., 2001; Pâques, 2004) crops is the development of genetic
and recently discussed by Miles (2007) in the variation. However, in populations with
frame of apomixis for cultivar development a high frequency of apomictic plants,
in tropical forage grasses. The proposed both facultative apomicts and completely
“evolutionary breeding approach” for Musa sexual plants can usually be found, and
spp. (Ortiz, 1997) is also in the narrow such genotypes can be used to develop
sense a reciprocal recurrent selection new genetic variation. For breeding, it is
scheme. However, subsequent application important to find or develop a system in
of reciprocal recurrent selection is rarely which both apomixis is maintained (variety
found in practice, although we think that development) and sexual reproduction is
this is the way ahead to exploit heterosis restored (population improvement) so as
and achieve more breeding progress in to be able to develop new genetic variation.
clonally propagated crops. This can be compared to male sterility
systems used in hybrid seed production. It no longer apply. Moreover, breeding TPS
is interesting that apomixis is distributed potato as a cross-fertilized crop will not lead
across many plant families. It appears not to to completely homogenous varieties. This
be controlled by a complex genetic system. is the reason why only a few TPS varieties
An example is Guinea grass (Panicum have been developed in the Northern
maximum), in which the sexual tetraploids Hemisphere. All these have been exclusively
are recessive homozygous (aaaa), whereas used for home garden production. However,
apomictic genotypes carry a dominant we think that by using hybrid selection
allele and are heterozygous (Aaaa) (Savidan, schemes and inbreeding in two separate
1983). So far, studies on apomixis have been gene pools it should be possible to develop
mainly made in tropical grasses, but more more and more homogenous and attractive
and more attention is being paid to rice and TPS varieties.
maize. There are opinions that apomixis The advantages of TPS are mainly
systems will become available to breeders, of interest in tropical and subtropical
and in this context gene isolation and an regions of the world. Under these climatic
‘apomixis gene’ have been mentioned conditions, the production, storage and
(Savidan, 2000). However, so far there is no transportation of potato planting material
such apomixis system usable in breeding is difficult. Moreover, potato yields are
programmes. The major problem is that considerable lower in tropical and
plants with the same genotype can express subtropical regions of the world than in
different degrees of apomixis. the Northern Hemisphere, so that about
20 percent of the harvest is needed as
13.9 PROPAGATION OF POTATOES BY planting material. Hence TPS varieties in
SEED the tropics can have 20 percent lower yields
Finally, the option that clonally propagated compared to clonally propagated potato
crops can be propagated by sexual seeds is varieties and remain competitive. About 20
considered. In many countries there have TPS varieties have been developed. Most
been research projects in which potatoes interesting are those varieties developed
were cultivated by seed. These are potatoes from recombination of the andigena and
that are sown instead of planted. Since the tuberosus gene pools. However, the original
planting material of clonally propagated idea of raising seedlings in nurseries and then
potatoes is often called a ‘seed’ potato, the planting seedlings into the field by hand has
term ‘true potato seed’ (TPS) was introduced. not been adopted. What has been adopted is
The use of TPS has two principle advantages: to raise TPS varieties in seedling nurseries to
the most important potato diseases cannot obtain healthy planting material, and then
be transmitted in true seed, and only a few to cultivate these TPS varieties for several
hundred grams of TPS are needed to cultivate growing seasons as a clonally propagated
a potato field, where usually several tonne crop, and to request true seed again after
of tubers are needed (Simmonds, 1997). yield declines are significant due to declining
This is associated with two disadvantages: health status (Fuglie, 2001). However, today,
potatoes grown from seed are weak in not more than 10 000 ha of TPS are grown,
vigour and are sensitive to many factors, mainly in Asia, which trace back to about
and the breeding method and advantages eight TPS varieties. The future of TPS is
of breeding clonally propagated crops can debatable. From the breeding perspective,
the future of TPS will mainly depend on the Americas (41 million tonne) and Africa
working with two gene pools, in which a (16 million tonne) (FAO, 2006). The top 20
certain extent of inbreeding is applied, with potato producing countries are China (22
subsequent use of general combining ability percent), The Russian Federation (12 percent),
to improve these two gene pools. India (8 percent), Ukraine (6 percent), United
States of America (6 percent), Poland (3
13.10 POTATO percent), Germany (3 percent), Belarus (3
Breeding potatoes has been reviewed by percent), Canada (2 percent), France (2
Tarn et al. (1992). The andigena potato percent), United Kingdom (2 percent),
(Solanum tuberosum subsp. andigena; Turkey (2 percent), Netherlands (1 percent),
autotetraploid with 48 chromosomes) Bangladesh (1 percent), Brazil (1 percent),
originated in the highlands of South Romania (1 percent), Peru (0.8 percent),
America about 5000 BC, while today two- Spain (0.6 percent), Nepal (0.5 percent) and
thirds of world potato production is in Pakistan (0.5 percent).
temperate latitudes. Following introduction
into Europe, andigena evolved into the Irish Breeding objectives
potato (S. tuberosum subsp. tuberosum), Characteristic of potato breeding is the
which is mainly characterized by day-length large number of breeding objectives. For
neutrality, uniformity of tuber shape, shorter the Irish potato, quality traits are at least
crop duration and higher harvest index than as important as yield. Moreover, breeding
andigena. Andigena remains the predominant for resistance against numerous pest and
cultivated potato in the Andes, whereas the diseases, e.g. numerous viruses (potato leaf-
Irish potato is the potato of commerce roll virus (PLRV), Potato virus Y (PVY)
in long-day temperate climates. Potatoes and Potato virus X (PVX)), Late blight
introduced into other, tropical, regions of (Phytophthora infestans), dry rots (Fusarium
the world trace back to breeding populations spp.), soft rot and blackleg (Erwinia spp.),
derived from crossings between andigena cyst-forming nematodes (Globodera
and Irish potato. However, in the Andean rostochiensis and G. pallida) have major
region, seven other potato species are still importance in long-day temperate as well
in cultivation; most important are phureja as in tropical temperate climates. For the
(S. phureja; diploid with 24 chromosomes), andigena potato, these pests and diseases
limeña, ajanhuri and rucki. In addition to are of nearly similar importance (i.e. late
these cultivated species, 160 wild potato blight can destroy the whole crop in cool,
species are known (Hawkes, 1979 and 1981; high-altitude regions, especially when the
Spooner and Hijmans, 2001), so that potato weather is wet).
might have the largest gene pool among crops. There are clear differences between
Wild and indigenous species are important tropical temperate and tropical hot
resources of pest and disease resistance for climates. At temperatures above 25°C,
andigena and Irish potato. The evolution of Late blight and cyst-forming nematodes
the potato was described in the introduction decline rapidly in importance, but Early
of this chapter. Asia and Europe are the blight (Alternaria solani) and root-knot
world’s largest potato producing regions, nematodes (Meloidogyne spp.) take
with annual production of about 130 and over, and Bacterial wilt (Pseudomonas
128 million tonne, respectively, followed by solanacearum) is widespread in tropical
lowlands. The phureja potato (for PVY and with considerable levels of total carotenoids,
Late blight) and the wild species S. acaule including pro-vitamin A.
(for PLRV, PVX and both Globodera spp.)
and S. demissum (for PLRV, PVY, and late Breeding methods
blight) are important resources in breeding Crossing is relatively easy. In nature,
for tolerance and resistance. It should be crossings occur easily by open pollination
noted that the pests and diseases presented by insects. For breeding purposes the
represent only the most important species. flower architecture of the potato allows
Ross (1985) provides a list of resistance easy emasculation and controlled hand
sources in wild potato species. pollination. A fruit with about 200 seeds
However, it is possible to find tolerance develops from each successful pollination. In
or resistance genes in cultivated and wild commercial breeding, controlled crossings
potatoes against nearly all potato diseases. are usually made (both parental genotypes
An exception is Bacterial wilt, for which so are clearly defined). Genotypes with good
far no useful tolerance or resistance have performance over all traits and with a certain
been found for breeding purposes. Today, degree of genetic distance are recombined.
all new Irish potato varieties contain one The value of a cross combination is usually
or more resistance genes from wild and determined in test-crosses with 100 to
other cultivated potato species. For the 200 seeds per combination. Occasionally
Northern Hemisphere, yield, crop duration, plant breeding text-books recommend
tuber size, shape and flesh colour, eye combining parents with complementary
depth, starch content, storability, cooking traits. However, many breeders find that
characteristics, taste and suitability for this results in potatoes breeding in a ‘wild’
mechanical harvesting, as well as processing segregation, so that finally only genotypes
characteristic for chips (crisps) and French can be selected with moderate performance
fries (chips) are the most important quality over all traits. Each year, breeders plant
breeding objectives. For tropical regions, 10 000 to 200 000 seeds, which trace back
yield, regional adaptability, crop duration, to 150 to 200 cross combinations. Crossings
storability, cooking characteristics, taste and are made with flower sprouts obtained from
nutritional quality are the most important cuttings of field plants, grown in greenhouses
quality breeding objectives. Outside of the in nutrient solutions. Frequencies of
Andes, crop duration is one of the most successful crosses differ tremendously
important traits (i.e. in south-west and between parental combinations and about
central Asia there is a requirement for one-eighth to one-quarter of all parental
potato varieties with less than 80 days combinations cannot be recombined due to
crop duration). Recently, focus has been no flowering, low pollen quality, no fruit
given to improve pro-vitamin A, iron and formation or genetic incompatibility. For an
zinc concentrations in tubers to alleviate overview of overcoming crossing barriers
micronutrient malnutrition in tropical in potatoes, the reader is referred to Jansky
regions (potatoes have comparatively high (2006).
iron and zinc concentrations). This has Pre-breeding crosses are important
resulted in a separate breeding programme when one requires to improve one or two
for phureja, which has the highest iron and traits in an enhanced breeding gene pool
zinc contents among potatoes, together (e.g. shorter crop duration). Pre-breeding
is usually made in two or three cycles include evaluation at several locations in

of recombination and selection in which replicated plots. However, the propagation
the desired traits are incorporated in a coefficient in potato is very low (≈ 10) so
genetic background that is more close to that it takes about eight to ten years before
the enhanced breeding gene pool. This is final selections enter the variety release and
generally done for resistances genes from dissemination stage.
a wild parent or exotic variety. It often Today there is little investment in TPS
involves an additional selection step at a in the original sense. However, selection
different polypoidy level. The re-synthesis of families that are to a certain degree
of tetraploids by mitotic duplication of homogenous in crop duration, tuber form
diploid genotypes (colchicine treatment and shape, with some genetic diversity in
of seed, axillary buds, tuber germs, tuber yield and specific adaptation, are of
leave explants or callus) is usually not interest. The reason is that these can be
recommended, because mitotic tetraploids disseminated as seeds and farmers have
have considerably lower yields than meiotic the option to exploit genetic variation for
tetraploids. Meiotic tetraploids occur specific adaptation in a PPB approach. PPB
naturally in crossings between tetraploid has been successfully applied in Bolivia in
and diploid potatoes (4x × 2x) due to early breeding stages (Gabriel and Torrez,
meiotic anomalies that result in unreduced 2000) and in later breeding stages (FVS) in
gametes (Rowe, 1967; Jacobsen, 1980). Ecuador (Bonierbale, pers. comm.).
Breeding potato for tropical regions of
the world aims mainly at improvement of 13.11 SWEET POTATO
four gene pools: (i) the andigena (A) gene Breeding sweet potatoes has been reviewed
pool for short-day high altitudes; (ii) the by Martin and Jones (1986), Laurie and
andigena × tuberosus (AT) gene pool for van den Berg (2002) and Grüneberg et
short- and long-day temperate regions, al. (2009). The sweet potato (Ipomoea
with emphasis on selection for Late blight batatas, Convolvulaceae, hexaploid with 90
tolerance and PVY and PLRV resistance; chromosomes) is also known as batata,
(iii) the tuberosus × tuberosus (TT) gene camote or yam (United States of America).
pool for short- and long-day warm regions, The crop was domesticated in tropical
with an emphasis on selection for short crop America about 6000 BC and reached the
duration; and the (iv) phureja gene pool (P), Pacific and south-east Asian islands naturally
with emphasis on nutritional quality. In or by early seafarers before Columbus. The
the A, AT and TT gene pools at CIP about number of wild species in the genus Ipomoea
60 parents are recombined by controlled is large (more than 500 species). However,
crossings to raise about 20 000 seedlings, no wild form of I. batatas has been found.
whereas in the P gene pool the number of It is assumed that I. batatas developed from
recombined parents is considerable lower an interspecific cross between a diploid and
(about 30 parents). The selections start in a tetraploid Ipomoea species in the I. trifida
seedling populations for both resistance complex. It is possible to re-synthesize new
and tuber formation. In three selection Ipomoea hexaploids by hybridization of
steps the material is reduced to about 300 diploid I. leucantha and tetraploid I. littoralis
clones, which are evaluated in 2-row plots (Nishiyami, Miyazaki and Sakamoto, 1975.).
with two replications. Later selection stages The Spanish introduced sweet potato in the
sixteenth century into the Philippines, whence adapted to salinity, drought and marginal
it spread to other islands and the east Asian soil conditions (Woolfe, 1992).
mainland. Portuguese seafarers introduced The crop has recently received more
the crop into Europe, Africa and India. Today interest due to the very high levels of
it is cultivated in 117 countries in all tropical pro-vitamin A (concentrations of up to
and subtropical regions of the world. Asia is 700 ppm DM) in OFSPs, and hence as
the world’s largest sweet potato producing a vehicle to reduce vitamin A deficiency
region, with about 107 million tonne of problems in the world (Huang, Tanudjaja
annual production, followed by Africa and and Lum, 1999; Low, 2007). We observed
the Americas, with approximately 15 and up to 1 200 ppm β-carotene on a DM
3 million tonne, respectively. The top 12 basis in clones with variety potential in
producing countries are China (80 percent), our breeding population ‘Jewel II’ (this
Nigeria (2.8 percent), Uganda (2.2 percent), corresponds to 30 mg β-carotene in 100 g
Indonesia (1.5 percent), Viet Nam (1.2 fresh sweet potato storage roots. A pre-
percent), United Republic of Tanzania (0.9 schooler needs 4.8 mg β-carotene per day,
percent), Japan (0.8 percent), India (0.8 and it merits discussion as to what extent
percent), Burundi (0.7 percent), Kenya (0.6 OFSP should be recommended as baby
percent), Rwanda (0.6 percent) and United and weaning food. Moreover, storage roots
States of America (0.6 percent). Further provide medium levels of iron and zinc
important sweet potato producing countries (Woolfe, 1992). Recent finding of about
are Angola, Argentina, Bangladesh, Brazil, 50 ppm DM iron and 40 ppm DM zinc in
Cuba, Egypt, Ethiopia, Haiti, Korea deep orange fleshed sweet potato storage
(Democratic Republic of), Madagascar, Peru, roots (Burgos and zum Felde, pers. comm.)
the Philippines and Papua New Guinea, merits further investigation.
with annual production between 0.3 and The stems and leaves can have spinach-
0.5 million tonne (FAO, 2006). Nearly half like taste and some varieties are used in
of the sweet potato produced in Asia is used China specifically as a green vegetable.
for animal feed, with the remainder primarily Stems and leaves have on DM basis about
used for human consumption, either as fresh four times more protein, iron and zinc than
or processed products. In Africa, the crop storage roots. It appears that stems and
is cultivated almost exclusively for fresh leaves must be cooked to reach an acceptable
consumption. iron bioavailability, but investigations into
Sweet potato is a perennial vine, propa- iron bioavailability of sweet potato tops is
gated by cuttings, and usually cultivated as very limited.
an annual crop. The planting distances in There is new demand for purple-fleshed
fields vary. In Africa, planting distances are sweet potato due to the health-promoting
usually 1 m between rows and 30 cm within effects of anti-oxidant anthocyanin sub-
rows. In China, recommended planting stances, and cell lines for a potentially
distances are 75 cm between rows and ongoing production for the food industry
20 cm within rows. The crop duration is have been established (Konczak, 2006).
very short (4 to 6 months) and the crop is However, much more important appears
even cultivated in northern China. It pro- to be the demand for non-sweet sweet
duces more edible energy per hectare per potatoes, but few genotypes are non-sweet
day than wheat, rice or cassava, and is well (Kays, 2006). There is a very large genetic
variation for DM, starch and sugars in the use of the crop as animal feed and
sweet potato, and a strong positive correla- folder. The breeding for high DM and
tion has been observed for DM and starch, extractable starch is relatively easy: the
whereas a strong negative correlation was target is a high starch yield per hectare.
found between sugars and DM and starch However, currently, in many regions of
(Grüneberg et al., 2009). This is nearly the world the price of sweet potato starch
ideal for the breeding target of a non-sweet currently cannot compete with the price of
high-DM sweet potato type, and we think cassava starch. Only in large regions where
that the development of non-sweet sweet the growing period is too short for cassava
potatoes should not be too difficult. within the cropping system (e.g. China) is
there an economic demand for sweet potato
Breeding objectives varieties for starch production. Breeding
FPB started very late for sweet potato. for biofuel production is in its initial stages,
One of the first breeding programmes was and so far variety recommendations for this
established at Louisiana State University in purpose are made on the basis of screening
the 1920s. Today there are several strong existing successful varieties. The breeding
national breeding programmes (e.g. China, of purple-fleshed sweet potatoes as a
Japan, South Africa, Uganda, United separate breeding programme is a relatively
States of America and Uruguay) and one new trend, and so far only carried out on
international breeding programme, at CIP a small scale in Japan, Indonesia and Peru.
(Peru). Four major breeding objectives can Future targets are the non-sweet sweet
be clustered: (i) breeding of OFSP for potato, and quick cooking features (cv.
consumption of storage roots and leaves; Quick Sweet) (Katayama et al., 2006), as
(ii) breeding for high DM and extractable well as suitability for processing into chips,
starch; (iii) breeding for biofuel production, puree, juice, weaning and baby food, and
which has started in China (Dai Fu Ma, bread on the basis of a wheat-sweet potato
pers. comm.); and (iv) breeding of purple- flour mixture (Woolfe, 1992); these trends
fleshed sweet potatoes for consumption. appear nearly exclusively in east Asia, and
In breeding for consumption, it has to be for recent developments the reader should
considered that people in different regions consult proceedings, such as Liu (2008).
have very different taste preferences; the Major constraints on high yields are
extremes are low DM content, moist mouth pests and diseases, especially Sweet potato
feel, very sweet taste and deep orange flesh chlorotic stunt virus (SPCSV) and the sweet
colour, versus high DM, bland, dry mouth potato weevils. The prevailing diseases and
feel, low sweet taste and white, yellow or insects affecting sweet potato vary from
orange flesh colour. In breeding for human region to region. There are about 35 bac-
consumption, focus is more on high DM terial and fungal diseases, more than 20
OFSP varieties with elevated iron and zinc viruses or virus-like agents, 20 nematodes
concentration and a dry and less-sweet and 20 insect species known to affect sweet
mouth taste. This breeding is hampered potato (Martin and Jones, 1986).
by a strong negative genetic correlation Currently there are only four important
between storage root DM and storage root pest and diseases: SPVD, Alternaria, sweet
pro-vitamin A, iron and zinc contents. The potato weevils and the root-knot nematode.
breeding for human consumption includes The most important virus is whitefly-
transmitted SPCSV, which often occurs in this reason, a transgenic approach using
co-infection with Sweet potato feathery Bt genes has received attention. Recent
mottle virus (SPFMV – aphid-transmitted). findings of compounds in the latex of the
Clear synergistic disease effects are seen storage root skin and the effect of these on
with SPFMV and SPCSV (the so-called weevils might of interest for breeding (P.C.
SPVD virus complex). Generally, all Stevenson, H. Muyinza, D. Hall and R.
varieties need a certain degree of tolerance Mwanga, unpubl.).
to SPVD, and there is genetic variation for
SPVD (Mwanga, Yencho and Moyer, 2002). Breeding methods
Very high tolerance or resistance is needed True seed set occurs easily in nature by cross
in eastern Africa. Currently, it is assumed pollination (by insects, mainly bees), and for
that SPFMV and all other sweet potato breeding purposes the flower architecture
viruses (except SPCSV) are not important, of sweet potato allows easy emasculation
because sweet potato has an effective virus and controlled hand pollination. A skilled
defence system, which is broken by SPCSV technician can make 200 crossings per day,
(I. Barker, pers. comm.). with a success rate of 25 percent. From
The major fungal disease in subtropical each successful cross, two or three true
America is Fusarium wilt (Fusarium seeds are obtained. Not all sweet potato
oxysporum f.sp. batatas) and in the African parents flower readily, but flowering can
highlands the main problem is Alternaria be easily induced by grafting on Ipomoea
storage root, leaf spot and stem blight nil (2n = 30 chromosomes). It should be
(Alternaria spp.). Although there are many noted that frequencies of successful crosses
bacterial and fungal diseases with a wide differ tremendously between parental
distribution, high levels of tolerance or combinations, and about one-third of all
resistance are frequently found. This is also parental combinations are incompatible,
true for resistance to nematodes. with no seed formation. The sweet potato
There has been recurrent success in seed has a hard coat and needs to be
breeding for root-knot resistance against scarified with concentrated sulphuric acid
new races of Meloidogyne spp. (Martin to obtain even and rapid germination. In
and Jones, 1986). However, in regions a well managed breeding nursery, after 3
with a pronounced dry season, the greatest months it is possible to obtain 40 to 60
constraints are sweet potato weevils (Cylas cuttings from a true seed plant if the plant
formicarius elegantulus in all parts of the is grown in the field, and 20 to 30 cuttings if
tropics, C. puncticollis and C. brunneus the plant is grown in a pot in a greenhouse.
in Africa, and Euscepes postfasciatus in The extreme genetic make up of the crop
the West Indies). It has been an objective (hexaploid, highly heterozygous, open-
to find weevil resistance for more than pollinated by insects with true seed set
50 years, but differences in weevil attack occurring easily), the short crop duration
probably depends on preference factors (4–5 months), and the rapid propagation
of the weevil. It is believed that dense (40 to 60 cuttings from one plant) permits
storage roots developed deep below the soil the design of a very efficient and rapid
surface are less susceptible than less dense, breeding system.
moist-fleshed storage roots. No effective The recombination of parents is still
weevil resistance has been found so far. For usually carried out in polycross nurseries
by open pollination. Polycrosses have 13.12 CASSAVA

been considered as very efficient in sweet Breeding cassava has been reviewed by
potato breeding (Martin and Jones, 1986). Byrne (1984), Bonierbale et al. (1994) and
However, theoretically controlled crosses Ceballos et al. (2004). Cassava (Manihot
must be more efficient, provided that high esculenta, Euphorbiacaea, diploid with 36
selection intensities can be reached, which chromosomes) originated in South America.
depends on technical skills and costs. Only The crop is also known as manioc and yucca.
a few breeding programmes are making Wild Manihot species—weedy sub-shrubs,
(at least to any major extent) controlled shrubs and trees—are principally found
crosses (e.g. in China, Mozambique, Peru in dry regions of Mesoamerica and South
and Uganda). The numbers of recombined America. The highest density of diversity
parents vary between 20 and 120, and the is found in west-central Brazil. Many
number of genotypes raised per population wild Manihot species show considerable
(true-seed plants) varies between 5 000 tuber production and it is assumed that
and 30 000. Selection of parents is almost M. esculenta was selected from one or
exclusively carried out on the parental several of these wild species in the northern
performance per se. In China, Uganda part of South America or in west-central
and at CIP in Peru, the information from Brazil. The crop was disseminated by tribal
progeny test crosses is used to repeat migrations and its variability increased by
good cross combinations on a larger scale selection for agronomically preferred types
(2 000 to 3 000 genotypes per cross). and further hybridization with wild species.
In recent years, CIP has established two Cassava was introduced in the fifteenth
genetically divergent populations to test century into West Africa by the Portuguese
heterosis and general combining ability in from Brazil, and from there it spread to
applied breeding material. There are plans eastern Africa, Madagascar and southern
to change from a selection of parents by India. Moreover, it was introduced in the
parental performance per se to a reciprocal sixteenth century into the Philippines by
recurrent selection scheme based on general Spanish traders from Mesoamerica. Today
combining ability. Selection of genotypes the crop is cultivated worldwide in lowland
for variety development is usually carried tropics. World production of cassava root
out as described in the section of the general was estimated to be about 226 million tonnes
breeding scheme for clonally-propagated in 2006, with most production in Africa,
crops. Starting with recombining parents, where 122 million tonnes were grown,
it takes on average 7 to 8 years until while 67 million tonnes were grown in Asia
variety release. At CIP, Peru, an accelerated and 37 million tonnes in Latin America and
breeding scheme is used in which temporal the Caribbean (FAO, 2006). The top ten
variation of test environments are replaced cassava producing countries are: Nigeria
by spatial variation of test environments. (18 percent of world production), Brazil (12
This accelerated breeding scheme takes on percent), Thailand (10 percent), Indonesia
average 3 to 4 years until variety release. It (9 percent), Democratic Republic of the
appears that there are funding opportunities Congo (8 percent), Ghana (5 percent),
to implement this breeding scheme in United Republic of Tanzania (4 percent),
Africa, particularly in Ghana, Uganda and India (4 percent), Mozambique (3 percent),
Mozambique. and Angola (3 percent).
Cassava adapts to a wide range of eco- a whitefly-transmitted virus widespread in

logical conditions and is known for its Africa and India; (ii) Cassava brown streak
tolerance of low soil fertility, drought and disease (CBSD); and (iii) Cassava bacterial
pests. The growing period is long, between blight (CBB), caused by Xanthomonas
7 and 18 months. The yields are very high campestris pv. manihotis, which can have
(about 30 to 40 t/ha under commercial devastating effects on yield in Africa. Of
practice). However, the protein content regional importance in Latin America and
of cassava is low (<3 percent DM), which the Caribbean is Frogskin disease, suspected
makes the crop ideal for starch produc- to be caused by a virus. Aside from these,
tion. Cassava is often grown in low input cassava is much less affected by disease than
production systems, particularly when it other tropical crops, the only other two
is grown as a food crop. Planting material of importance being Cassava anthracnose
is easily obtained from plant stems avail- disease (Colletotrichum manihotis) and
able from the farmers’ own or neighbour- root rots (Phytophthora drechsleri and
ing fields. About 70 percent of cassava is Rhizoctonia spp.) (CIAT 2001; Hillocks
grown by small-scale producers for direct and Wydra, 2002).
human consumption. The crop tolerates The major pests of cassava are
more drought, lower soil levels of nitrogen, nematodes (Meloidogyne spp.), whiteflies
potassium and phosphorus, lower pH and as a vector of CMD, Cassava green mites
higher aluminium levels than most other (Mononychellus spp. and Tetranychus spp.),
crops. Under these conditions, yields are cassava mealybug (Phenacoccus spp.), and
about 7–10 t/ha. Cassava is often found the grasshopper (Zonocerus elegans). Pests
in mixed stands, together with a variety of and diseases, together with poor cultural
other food or cash crops. Estimates indicate practices, combine to cause yield losses as
that at least one-third of the cassava grown high as 50 percent. In the late 1980s, a new
worldwide is intercropped (Cock, 1985). strain of CMD occurred in Uganda that
made the virus more harmful. This mutated
Breeding objectives virus has been spreading and is now found
FPB started in isolated programmes in throughout Uganda, Burundi, Cameroon,
the early 1900s when cultivation was the Democratic Republic of Congo and
extended by several colonial governments Rwanda (Thresh and Cooter, 2005). Next in
as a safeguard against famine, and breeding importance in breeding are more short and
new clones with resistance against cas- thick storage roots with high starch content.
sava mosaic disease (CMD) was required. This is important for mechanical harvesting,
Cassava breeding programmes started in but makes also manual harvesting easier. It is
Brazil (in the 1930s), India (in the 1940s), desirable for the roots to be as far as possible
Indonesia (in the 1950s) and at two inter- horizontal in the soil and near to the soil
national institutions: CIAT, Colombia, surface. Breeding selects for plants with lower
(in the 1970s) and IITA, Nigeria, (in the height and higher harvest index. In cassava
1970s). These institutions have developed a breeding for human consumption, the focus
very successful cassava breeding network. is on yield and quality such as low fibre, low
In cassava breeding, three diseases have levels of cyanogenic glucosides, high protein,
been the highest priority for decades: elevated pro-vitamin A, iron and zinc
(i) Cassava mosaic disease (CMD), which is concentration in the storage roots, reduced
post-harvest physiological deterioration and depression and wide segregation in cross

regional preferences for the peel of the roots progenies. Time of flowering depends on the
(Ceballos et al., 2004). Cassava varieties are genotype. There are types in which flowering
often categorized as either ‘sweet’ (actually starts about two months after planting, as
‘not bitter’) or ‘bitter’, signifying the absence well as types that do not flower until after
or presence of toxic levels of cyanogenic 24 months or more. This makes planned
glucosides. The so-called ‘sweet’ cultivars recombination difficult. Earlier and more
can produce as little as 20 mg/kg cyanide in abundant flowering is obtained by foliar
fresh roots, while ‘bitter’ ones may produce application of indole acetic acid (IAA) and
more than 50 times as much. Additionally, naphthalene acetic acid (NAA). The female
an important breeding objective is to flowers are large, nearly always located at
develop more clones with high adaptation the base of the inflorescence, and open first.
to drought-prone environments. The genetic The female flowers normally open 10–14
variation in cassava for pro-vitamin A days before the males on the same branch,
concentrations is small. However, breeding but self-fertilization can occur because male
for yellow cassava genotypes with a pro- and female flowers on different plants of the
vitamin A concentration of 15 ppm appears same genotype can open simultaneously.
to be possible. Additionally, a transgenic The proportions of self- and cross-
approach is used to introduce the β-carotene pollinated seed produced depends on
pathway into cassava (J. Tohme, pers. comm.). genotype, planting design and the type
Breeding for commercial production also of pollinating insects present (5 percent
selects for plants with shorter height and self-pollination occurs naturally). Both
higher harvest index – giving more stability the stigma and the pollen are sticky and
and resistance against storms – and extensive pollination is easily carried out by honey
branch formation, quickly forming a full bees. In the Northern Hemisphere, cassava
canopy of leaves not too close to the soil usually flowers from July to January, with a
(Byrne, 1984). peak between September and November. In
the Southern Hemisphere, it usually flowers
Breeding methods from January to July, with a peak between
Cassava is a monoecious, highly heterozygous March and May. Tall plants with less
plant. All 36 chromosomes show regular branching are less floriferous than highly
bivalent pairing at meiosis. However, in branched, low growing ones. To make
both cassava and Manihot glaziovii (sect. a controlled cross between two parents,
Arboreae) there is evidence of polyploidy unopened flowers are first enclosed in
from studies of pachytene karyology. There muslin bags and the chosen pollen applied
are three nucleolar chromosomes, which is to the stigmas as soon as the female flowers
high for true diploids, and duplication for open. The muslin bags are then replaced
some of the chromosomes. This indicates with netting bags to catch the seed when the
that Manihot species are probably segmental ripe fruits dehisce explosively. The fertility
allotetraploids derived from crossing between of clones is variable and can be very low;
two taxa whose haploid complements had an average of one or two seeds per fruit
six chromosomes in common but differed is common in controlled pollination. Seed
in the other three (Nasser, 2000). Cassava matures 70 to 90 days after pollination. The
shows self-fertility with strong inbreeding fruits are collected when the coat begins to
shrivel and are sun dried until they shatter, ca. 100 clones are tested in replicated trials
releasing hybrid seeds that are ready for at three locations, and consumer acceptance
germination. Cassava seed have a very short is assessed. Final selections are multiplied
dormant period and germinate quickly. and enter dissemination in year 6.
No scarification is necessary. Few seeds In eastern and southern Africa, 10 000 to
germinate unless the mean temperature 50 000 true-seed plants are raised for the first
exceeds 24°C, with a temperature exceeding selection step and screened for resistance
30°C for at least part of the day; the best rates to major diseases and pests at 1, 3, 6 and 9
occur at 30–35°C. A dry heat treatment of months after sowing, namely East African
14 days at 60°C is also beneficial for newly cassava mosaic disease (EACMD), African
harvested seeds. If temperatures permit and cassava mosaic disease (ACMD), Cassava
irrigation is available, the easiest method is brown streak disease (CBSD) and CBB.
to sow the seeds direct into the soil. This In a second selection step, 2 000 clones are
is successful at IITA because temperatures planted in single-row plots (3 to 5 plants) at
from January to March range from 30° 1×1 m spacing. The observations made in the
to 35°C. At CIAT, seeds are frequently first year are repeated again at 1, 3, 6 and 9
planted in a screen house and the emerging months after planting. Each clone is scored
seedlings held until they reach 20–25 cm for yield, and agronomic characteristics
before being transplanted to well prepared assessed, such as branching height and angles,
soil with good moisture conditions. canopy and number of stems per plant. In a
Since many national programmes do third selection step, 20 to 50 clones are
not have a continuous cassava crossing grown in preliminary yield trials in single
programme, they rely on distribution of pre- rows with ten plants per clone and three
selected clones from the two international replications at one to three locations. In year
institutions, CIAT and IITA. The improved 6, the final selections are taken on-farm and
germplasm generated is distributed either into national variety release trails.
in the form of elite genotypes transferred In the Americas and Asia, cassava
in vitro, or as populations of recombinant improvement is closely linked with the insti-
seeds (full-sibs or half-sibs). Cassava tutions Embrapa (Brazil), FCRI Rayoung
breeding operates with larger populations (Thailand) and CIAT (Colombia). In con-
than potato or sweet potato. trast with Africa, there are no extremely
In West Africa, up to 100 000 true devastating diseases. CIAT established
seed plants are raised from field-sown 50 000 seedling selections for particular
seed, which are screened in a first selection climatic zones. Up to 20 parents from each
step for resistances to CMD and CBB. gene pool are disseminated for evaluations
At harvest, selection is for compact roots to national centres in similar edapho-cli-
with short necks, stems branching at about matic zones. From this programme a very
100 cm, with low HCN in the leaves. broad range of improved diversity has been
In the second selection step, about 3 000 developed and distributed worldwide.
clones are grown in small, non-replicated Generally, MVs in Asia can be traced back
plots. Further selection is made for disease to 100 crosses between Asian and American
resistances, yield potential and root DM parents (Kawano, 2003). Recent findings
content, and the HCN in the roots is assayed show that the general combining ability for
enzymatically. For the third selection step, cassava fresh root yields are clearly larger
than the specific combining ability across (Simmonds, 1976; Ortiz, 1995). They are a
contrasting environments (Ceballos et al., staple in Samoa, the Philippines, south India
2004). This is a clear indication that heterotic and the West Indies. Around 87 percent of all
gene pools in cassava can be formed and bananas and plantains grown worldwide are
exploited by improving two gene pools with produced by small-scale farmers for home
a reciprocal recurrent selection scheme. consumption or for sale in local markets.
About two-thirds of world production is
13.13 BANANA OR PLANTAIN dessert bananas and one-third plantains.
Breeding bananas and plantains has The fruit export market comprises only
been reviewed by Rowe (1984), and Jain one-sixth of total world production. The
and Swennen (2001) have edited recent banana is the number one fruit crop in
proceedings on banana improvement, with the world, with about 70.5 million tonne
a main emphasis on biotechnology. Banana produced annually. The top ten producing
and plantain (Musa × paradisiaca, Musaceae, countries are India (24 percent), Ecuador (9
usually triploid with 33 chromosomes) percent), Brazil (9 percent), The Philippines
originated in Southeast Asia. The term (8 percent), China (8 percent), Indonesia (5
plantain is used for those bananas that are percent), Costa Rica (3 percent), Mexico (2
palatable only when cooked. The crop was percent), Thailand (2 percent) and Colombia
introduced into Africa about 3 000 BPE. (2 percent). Plantains are grown as a staple
Introduction into the Americas came after food in 52 countries worldwide with a total
1 500 AD. Today the crop is cultivated production of 34 million tonne. The top ten
worldwide in the tropics. Bananas and plantain producing countries are Uganda
plantains evolved from two diploid (30 percent), Colombia (9 percent), Rwanda
wild species, Musa acuminata (AA) and (8 percent), Ghana (7 percent), Nigeria (6
M. balbisiana (BB) in the Eumusa series percent), Peru (5 percent), Cote d’Ivoire (4
(x = 11) of the genus Musa. An exception percent), Congo (4 percent) and Kenya (3
is the small group of ‘Fehi ’ bananas in percent) (FAO, 2006).
the Pacific, which have their origin in the Bananas and plantains are one of the
Australimus series (x = 10) of Musa. All very few crops in which breeders are still
export fruit bananas are triploids (AAA) trying to find an appropriate conventional
and originated from M. acuminata. All breeding method to develop new MVs.
plantains and several locally preferred fruit Nearly all cultivars are FVs and have been
bananas are hybrids between M. acuminata selected from genetic variation developed
(AA) and M. balbisiana (BB). The higher by natural evolution. In cases of crop failure
dry matter (about 5–8 percent) and higher due to new pathogens and diseases, FPB still
starch content of plantains compared to focuses on identifying alternative cultivars
pure M. acuminata cultivars is attributed within existing genetic variation (collections
to the BB genome. The AAB cultivars and large screening programmes). Hence,
have long curved fruits and appear like an important source for identifying ‘new’
an oversized export banana. They are cultivars are germplasm collections held in
important food crops in south India, eastern trust in genebanks, such as the International
and central Africa and tropical America. Musa Germplasm Collection in Leuven,
The ABB cultivars have thick straight fruits, Belgium. Spontaneous mutants in Musa have
which are much shorter than the AAB types played a very important role in banana and
plantain breeding, including the replacement widely distributed. However, for several
of the export banana cultivar ‘Gros Michel’ of these FHIA cultivars, taste and cooking
(susceptible to Panama disease or Fusarium qualities are still problematic (Roux, pers.
wilt (Fusarium oxysporum sp. cubense) by comm.). Further breeding programmes
‘Cavendish’ banana cultivars, which are have been set up at the Empresa Brasiliera
resistant to most fusarium wilt pathogens, de Pesquisa Agropecuaria (Embrapa) in
and the replacement of the plantain cultivar Brazil, the Instituto de Investigaciones
‘Horn plantain’ (AAB) (susceptible to Black en Viandas Tropicales (INIVIT) in Cuba,
sigatoka (Mycosphaerella fijiensis)) by the the Centre Africain de Recherches sur
‘Laknau’ cultivar (AAB), which is tolerant Bananiers et Plantains (CARBAP) in
to Black sigatoka and closely resembles the Cameroon, the International Institute of
Horn plantain (Stover, 1972). However, the Tropical Agriculture (IITA) in Nigeria, and
cooking qualities of Laknau are inferior the National Research Centre on Banana
to Horn plantain. Owing to the low level (NRCB) in India.
of occurrence of spontaneous mutations,
mutagenic agents and mutation breeding Breeding objectives
have often been used to generate new In breeding, resistance against Panama
genetic variation in bananas and plantains, (Fusarium wilt) and sigatoka diseases are
followed by screening programmes for in the foreground. In the first half of the
plants with resistance or tolerance to twentieth century, Panama disease destroyed
pest and diseases, coupled with desirable approximately 40 000 ha of bananas in
agronomic qualities (e.g. tolerance to Central and South America. Fortunately,
Panama disease; tolerance to the toxin of resistant Cavendish cultivars could substitute
Mycosphaerella fijiensis; short; larger fruit for the predominantly grown Gros Michel
size; and earliness). The FPB programmes variety. However, Cavendish cultivars are
for bananas and plantains started in the not resistant to all fusarium wilt pathogens
early 1900s, to develop new AAA cultivars (i.e. race 4). It should be noted that Panama
for the export market, with resistance disease cannot be controlled chemically,
against Panama disease or Fusarium wilt so that use of resistant varieties is the only
(Fusarium oxysporum f.sp. cubense). way to maintain production in regions with
Despite continued breeding efforts, no new challenge from this disease. The leaf spot
banana and plantain cultivar acceptable diseases caused by Mycosphaerella musicola
by farmers and consumers was bred until (Yellow sigatoka) and M. fijiensis (Black
the 1980s (Roux, 2001). Nevertheless, by sigatoka) are costly pathogens and must be
the end of the twentieth century, efforts regularly controlled by fungicides. Cultivars
to improve Musa started to focus on the with an AAA genome are very susceptible
use of diploid and tetraploid gene pools to both sigatoka diseases. The Horn
to develop triploid and tetraploid bananas plantain is resistant to Yellow sigatoka, but
and plantains. To date, the first improved susceptible to Black sigatoka. The latter
cultivars (AAA, AAAA, AAB, AAAB and disease threatens continued cultivation of
AABB), developed at Fundación Hondureña the plantain food crop. Triploid cooking
de Investigación Agrícola (FHIA) in bananas of the ABB type, such as ‘Chato’,
Honduras through the International ‘Pelipita’ and ‘Saba’, are highly tolerant
Musa Testing Program (IMTP), have been to the Black sigatoka pathogen. However,
Chato is susceptible to bacterial wilt or tillering capacity (Ferwerda and Wit, 1969;
Moko disease caused by Pseudomonas Rowe and Richardson, 1975; Persley and
solanacearum and to race 2 of Fusarium De Langhe, 1987).
oxysporum f. sp. cubense, while Pelipita does
not meet flavour and fruit-shape preferences, Breeding methods
so that currently only Saba remains as a Triploid bananas and plantains are vegetatively
possible substitute for the Horn plantain. parthenocarpic, i.e. no pollination is
Moreover, nematodes, mainly the burrowing necessary for fruit development. In diploids,
nematode (Radopholus similis), are a major pollination often results in seeded fruits.
constraint to bananas in monoculture, Diploids are not suitable as varieties since
and outside of the Americas the Bunchy fruit size and plant vigour are low. However,
top virus is widely distributed, which is diploids are the basis for crop improvement.
transmitted by the banana aphid (Pentalonia In the initial stages of breeding efforts, a few
nigronervosa). Many diploid accessions of seeds per bunch in some triploid varieties
M. acuminata subsp. malaccensis and M. a. were used when these had been pollinated
subsp. burmannica are resistant to races by diploid genotypes. The reason for this
1, 2 and 4 of Panama disease. Sources of seed production and genetic variation is the
resistance to Yellow sigatoka are available in formation of unreduced triploid gametes in
several subspecies of M. acuminata, while some triploid female parents after pollination
M. a. subsp. burmannica is highly tolerant within diploid male parents, which produces
to the Black sigatoka fungus. The tolerance reduced haploid gametes. The progenies of
in M. acuminata accessions to sigatoka these crosses are tetraploid. This method
diseases is apparently controlled by several was used to generate genetic variation with
dominant genes. Resistance to the burrowing the female banana parent Gros Michel and
nematode has been found in the ‘Pisang Jari the female plantain parent Laknau (AAB),
Buaya’ group of diploid accessions. The which closely resembles the Horn plantain.
resistance is controlled by one or very few Tetraploid hybrids (AAAA) from crosses
dominant genes and has been incorporated with Gros Michel were resistant to Panama
into diploid and polyploid progenies. Today, disease and closely approached commercial
several FHIA varieties are resistant to acceptability, but the inferior agronomic
burrowing nematodes (Kalorizou, Gowen characteristics of the diploid parents
and Wheeler, 2007). were also present in the hybrids. Triploid
Among agronomic qualities, dwarfness hybrids derived from crosses between these
is most important in bananas and plantains, tetraploid hybrids and diploid genotypes
because they are often grown in areas were useless. Unfortunately, the cooking
with periodic strong winds. Dwarf and qualities of hybrids derived from Laknau
semi-dwarf mutants have been found in were also inferior to those of the Horn
many diploid and triploid bananas and plantain. No seeds have been produced
plantains. Examples are ‘Highgate’ (a dwarf from Cavendish clones and no other
mutant of Gros Michel) and the Cavendish suitable triploid parents—except Gros
cultivar ‘Grand Nain’. In dwarf diploids, Michel and Laknau—for seed production
the dwarfness character is controlled by unreduced gametes have been found.
by a single dominant gene. After this in This breeding method has not succeeded in
importance are fruit characteristics and creating acceptable new varieties. However,
the major finding of this work was that it Thompson and Aked, 2000; Ludger, 2005;
is necessary to improve the diploid male Kalorizou, Gowen and Wheeler, 2007).
parent gene pool to increase the chances However, to our knowledge, no PPB has
of developing either new tetraploid or new been applied in early breeding stages. Most
triploid varieties. likely the reason for this is that almost
Today, banana and plantain breeding no diploid clone in its performance per se
aims at producing tetraploid and triploid would achieve acceptability by farmers.
varieties on the basis of diploid accessions Nevertheless, the future of banana and
resistant to various diseases, and the plantain breeding, as in other clonally
continuous improvement of this diploid propagated crops, should be seen in testing
gene pool for agronomic qualities (i.e. plant the combining ability between two gene
height, fruit characteristics and tillering pools and in the improvement of two
capacity) as well as high pollen production. gene pools on the basis of the general
Crossings within the diploid gene pool are combining ability and reciprocal recurrent
complex: the diploid ‘SH-2095’, which was selection. In banana and plantain breeding,
later successfully used in tetraploid variety such a breeding system can be established
development, was derived from a four-way by a seed-fertile diploid gene pool with
cross of three diploid cultivars and one wild high pollen production and a seed-fertile
accession ((‘Sinwobogi’ × ‘Tjau Lagada’) × tetraploid gene pool, which is used as the
(‘Guyod’ × a wild Musa acuminata subsp. male parent. In such a breeding programme,
malaccensis)). Nevertheless, the genetic basis PPB could easily be incorporated. However,
of diploid pollen parents with improved the important information provided by
agronomic performance is considerably the farmers would not be seen in the
wider than in the past. The currently evaluation of clone performance per se in
best diploids are continually crossed on the diploid and tetraploid gene pool, but in
triploid Highgate and Laknau, which the numbers of acceptable clones per cross
produces unreduced triploid gametes, as combination and family between genotypes
well as on seed-fertile tetraploids with good of the diploid and tetraploid gene pool, as
agronomic performance. The first results in described above in the section on selection
new potentially tetraploid varieties and the of parents and cross prediction.
later in new potentially triploid varieties.
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323
CHAPTER 14
Breeding for quantitative variables.

Part 1: Farmers’ and scientists’
knowledge and practice in variety
choice and plant selection
Daniela Soleri and David A. Cleveland

14.1 INTRODUCTION to farmers’ varieties (FVs), which include

This chapter focuses on the knowledge and landraces, traditional (folk) varieties selected
goals for selection of Third World farmers by farmers, modern varieties (MVs) adapted
in comparison with those of formal plant to farmers’ environments by farmer and
breeders. By Third World farmers (here- natural selection, and progeny from crosses
after simply ‘farmers’) we mean those in between landraces and MVs (sometimes
the relatively marginal (high stress, high referred to as creolized or degenerated MVs)
spatial and temporal variability) growing (Berg, 2009; Cleveland, Soleri and Smith,
environments of small-scale, traditionally- 1994; FAO, 1996; Zeven, 1998). Sustaining
based agricultural systems (hereafter simply and increasing crop production is essential
‘small-scale‘ or ‘Third World’ agriculture). for the survival of SSTW agriculture, and,
The assumption of conventional economic in this, seed saving and plant breeding have
development has for decades been that these critical roles to play
farmers would soon be absorbed into the In this chapter we review theory and
industrial sector, and food production would data on selection by farmers, and compare
shift to large-scale, industrial farms, and this it with selection by formal, scientific plant
scenario is still seen as desirable by many breeders (hereafter simply ‘plant breeders’
(e.g. Conway, 2003). There is, however, or ‘breeders’). Because selection by farmers
evidence that this small-scale Third World and formal plant breeders is based on
(SSTW) agriculture remains necessary for the same basic biological principles, their
feeding a significant proportion of the world understanding and practice of selection may
population, and will probably be necessary be similar. However, there are differences
in the future, even with production increases between farmers and breeders in the genotypes
in large-scale, industrial agriculture (Hazell and environments they work with, including
et al., 2007). More than 2 billion people live the types of agricultural systems for which
on almost 500 million small-scale farms (<2 they are selecting, as well as differences in
ha) in the Third World, including half of their experiences, technologies and goals
the world’s undernourished people and the for selection. Similarities and differences
majority of people living in absolute poverty in selection among farmers and among
(Nagayets, 2005). Economic re-structuring formal plant breeders also exist, for the same
beginning in the 1980s removed government reasons. Our goal in this chapter is to review
support for SSTW agriculture and led to what we know about these similarities and
migration from rural to urban areas, creating differences, and why understanding them
a crisis there (Hazell et al., 2007; Narayanan is important for collaboration between
and Gulati, 2002; Wise, 2007). In addition farmers and breeders to improve selection
to irreplaceable food production, SSTW for varieties that could help SSTW farmers
agriculture has other benefits: it operates in survive and prosper in the future.
many of the world’s centres of crop genetic We believe that respect for farmers and
diversity, where farmers conserve diversity their knowledge is essential for achieving
in the form of crop genetic resources in the maximum benefits from collaborative
situ, along with rich cultural and linguistic plant breeding. The greatest single mistake
traditions (FAO, 1996; Harlan, 1992). Plant plant breeders and other outside scientists
or crop genetic resources comprise wild can make is to assume they understand local
and weedy relatives of crops in addition agricultural systems. Even if their hypothe-
Breeding for quantitative variables. Part 1: Farmers’ and scientists’ knowledge and practice in variety
choice and plant selection 325
ses are accepted through local research, new on definitions: a farmer’s variety of a self-
details and perspectives are sure to arise, pollinated crop (e.g. of barley or rice) may
and it is only by having an open minded, be composed of diverse genotypes that,
respectful attitude that outsiders can hope from a plant breeder’s perspective, may be
to learn and reap the benefits of collabora- different varieties. Therefore discriminating
tion with local farmers. Such an attitude among these genotypes would be selection
facilitates new insights and understandings from the perspective of farmers as it can
that can improve the accuracy and relevan- change the genetic make up of their variety,
cy of the scientific work. If plant breeders but would be choice from the perspective
think of their interactions with farmers as of plant breeders as it would not change
tests of how complete or accurate farmers’ the genetic make-up of varieties as they
knowledge is, breeders will lose a critical define them. At a more fundamental level,
opportunity for supporting collaboration, farmers’ choice of populations and varie-
respect and collegiality, and for improving ties determines the diversity available for
the quality of their own work. Thus, at least hybridization and subsequent selection of
initially, experiments and discussions with plants. For all of these reasons, we can say
farmers should be seen as opportunities to that selection and choice together deter-
learn, not to teach. mine the degree to which varieties stay
the same, change between generations, or
14.1.1 Choice as distinct from selection evolve over generations.
It is important to differentiate between Farmers and plant breeders make choices
choice of populations or varieties that between varieties and populations, especially
does not change the genetic make-up of in the initial stages of the selection process
these units, and the selection of plants when choosing germplasm for making
from within populations or varieties, with crosses (for plant breeders), and in the final
the potential to change the genetic make- stages when choosing among populations
up of these units, which may eventually or varieties generated from those crosses
result in new varieties (Cleveland, Soleri for further testing (Hallauer and Miranda,
and Smith, 2000). Farmer criteria for both 1988: 159), or for planting (farmers) or
choice and selection include agronomic, release (plant breeders). Farmers’ choices
economic, culinary and aesthetic charac- of varieties or populations when saving
teristics, as well as minimizing perceived seed for planting, in seed procurement and
risk. While the distinction is commonly in allocating different varieties to different
made in some participatory plant breeding growing environments also affects the
literature (e.g. Witcombe et al., 1996), the genetic diversity of their crop repertoires,
terms ‘choice’ and ‘selection’ are often not and establishes the diversity on which future
explicitly defined, and in some writing may selection will be based. (For simplicity, in
be used interchangeably. Obviously, distin- the discussion of choice we will use the
guishing between these is partly a function term ‘variety’ to refer to both populations
of scale, as is most clearly seen in the case and varieties.)
of vegetatively propagated crops, in which
a single clone may be chosen to establish 14.1.2 A taxonomy of farmer selection
a new variety (e.g. with cassava; Pujol, A taxonomy of selection and its biological
David and McKey, 2005). It also depends effects can help to clarify the differences
FIGURE 14.1
Phenotypic selection classified according to the agent of selection, and farmers’ goals
when the farmers are the agents, applied to traditional agricultural systems
Phenotypic selection: identification of individual plants within a population that will

contribute genetic material to next generation (in each of the cases below, phenotypic
selection can have range of outcomes in terms of S, R and E).
Natural phenotypic selection by Artificial phenotypic selection by

biotic and abiotic factors in the farmers
environment not controlled by farmers,
e.g. climate, soil texture, pathogens,
pests
Direct phenotypic selection of plants

by farmers
Indirect phenotypic selection by
biotic and abiotic factors in the
environment (fields, store rooms)
managed by farmers, e.g. soil moisture
due to irrigation, intercropped plants of Intentional phenotypic selection.
other species, seed storage methods Farmers have explicit, conscious
selection goals
Unintentional phenotypic selection.

Farmers have no conscious goals for
phenotypic selection aside from Intentional phenotypic
obtaining seed; goals may be selection for R. Farmers have
unconscious, e.g. selecting large seeds conscious goals for eliminating
because they are easier to handle or changes in key varietal traits,
don’t fall through a basket; saving fruit i.e. inter-generational
from earliest producing plants for seed population maintenance
Intentional phenotypic selection for Intentional phenotypic selection for

goals other than R or E. Farmers have E. Farmers have conscious goals for
conscious goals for physiological, multi-generational population change to
morphological or phenological traits create new genotypes, e.g. by selection
like large seed size in Oaxaca, but not for seed color or plant structure
goals of intergenerational maintenance
or change
See text for key to symbols.

Copyright © 2008, D. Soleri, D.A. Cleveland, used with permission.
and similarities between plant breeders a bridge between these is useful (Soleri
and farmers. Selection can be categorized and Cleveland, 2005). For plant breeding,
according to the agent carrying out the most fundamental model of the
phenotypic selection, and the intention of relationship among phenotype, genotype
the agent when it is a human (Figure 14.1). and environment is assumed to be a good
While all types of selection function in model of reality that is the basis for PBK; we
both farmer and professional breeding, will assume it is also the basis for FK. This
professional plant breeders see intentional model is universally accepted by biologists,
phenotypic selection for micro-evolution including plant breeders, but they disagree
over generations (E) as the primary among themselves about its interpretation at
goal, with other types of selection either higher levels of generalization, for example
eliminated (e.g. applying irrigation to whether selection in optimal or marginal
eliminate drought selection), controlled for environments leads to genotypes that are
(e.g. in experimental plot design to reduce better adapted to marginal environments
E2 ), or used to optimize selection for (Ceccarelli and Grando, 2002) (see
E (e.g. roguing off-types) (Cleveland and Chapter 2). This variation in scientists’
Soleri, 2007). interpretations suggests that, if farmers
Figure 14.1 focuses on selection under do in fact think in terms of this basic
farmer conditions. Natural selection is not biological model, it would be a valuable
influenced by farmers, in contrast with comparator, facilitating understanding of
human or artificial selection. Artificial selec- variations (differences in higher levels of its
tion is both indirect, a result of the envi- interpretation) within and between FK and
ronments created by farmers and plant PBK on equal grounds.
breeders, e.g. in their fields and store rooms, We use the two parts of the model on
and direct, a result of human selection of which plant breeding is based (Cleveland,
planting material. Direct artificial selection Soleri, and Smith, 2000), as presented in
can be both unconscious or unintentional standard texts (e.g. Falconer and Mackay,
(based on implicit or correlated criteria), 1996: 189; Simmonds and Smartt,
when no conscious decision is made about 1999: 193).
the trait selected for, and conscious or inten- 1. Variation in population phenotype
tional (based on explicit criteria), the result (observable characteristics) ( P2 ) on
of decisions to select for certain traits. which choice (discrimination between
different groups of plants) and selection
14.1.3 A biological model to compare (discrimination among individual plants
farmer and plant breeder knowledge within a group) are based is deter-
and practice mined by genetic variation ( G2 ), envi-
Many plant breeders and other outsiders ronmental variation ( E2 ), and variation
who work with farmers make the mistake of in genotype (genetic constitution)-by-
assuming that western scientific knowledge environment (G×E) interaction ( GE2 ),
and practice is always more accurate and thus P2 G2 E2 GE
2
.
‘better’ than that of farmers. To have a 2. Response to selection (R) for a trait is
way of comparing plant breeder knowledge the difference between the mean of the
(PBK) and farmer knowledge (FK), a whole population from which the parents
neutral comparator that can function as were selected and the mean in the next
generation produced by planting those Soleri, 2002b). When FK differs from that
selected seeds under the same conditions. presumed by plant breeders’ interpretation
R is the product of two factors, h2 and of the model, we should try to understand
S (R = h2S), where S is the selection the difference in terms of the specific geno-
differential, the difference between the types and environments each works with,
mean of the selected parental group and as well as other factors in their experience.
the mean of the entire original population
(Allard, 1999: 101–102; Falconer and 14.1.4 Methods for understanding
Mackay, 1996: 189; Simmonds and Smartt, farmers’ knowledge and practice
1999: 193). Narrow sense heritability The best starting place for collaboration may
(h2) (that part of P2 that can be passed be simple interviews with a representative
directly from parent to progeny, the random sample of households. Such
additive variance, A2 ) = A2 / P2 . Thus, interviews can provide insights critical for
artificial phenotypic selection per se is collaboration. There are many resources
a process of identifying the individuals available describing how to conduct such
with specific phenotypic traits within a interviews (e.g. Cleveland and Soleri, 1991)
population that will contribute genetic and analyse them (e.g. Stern et al., 2004).
material to the next generation, and is The key requirements are that: (i) the
distinct from the heritability of those sample is representative of the human
phenotypic traits (see Section 14.5). population with which you are working,
In our use of the basic biological model, possibly requiring a stratified sampling
we make several assumptions. (1) It models approach, based for example on gender of
empirically observable patterns in the real farmers, household socio-economic status,
world. (2) Among both farmers and plant or dominant soil type on farms; (ii) people
breeders and other scientists, there are conducting the interviews are consistent,
some who are particularly good observers respectful, open and primarily listen
of their environments, crops and interac- to and document farmers’ answers and
tions between these if they occur, while comments; and (iii) questions are relevant
others are poor observers, resulting in for understanding and collaboration.
variation within groups. (3) Variation in In addition to simple questions to elicit
knowledge within and between groups can basic descriptive data (household size,
also be caused by experiences with different number working in farming, area sown to
genotypes and environments, and by dif- each crop, sources of planting seed, yields,
ferent values and pre-existing knowledge. etc.), methods such as scenarios and ranking
(4) Differences between FK or PBK and exercises may use hypothetical varieties
the model do not mean that either form to better understand farmers’ theoretical
of knowledge is wrong, and differences knowledge, or actual varieties they are familiar
between FK and PBK do not mean that with for insights into specific experiences and
either is inferior to the other. observations (Crossa, Bellon and Franco,
Thus, experiences under diverse circum- 2002; Soleri and Cleveland, 2005). For
stances can result in local interpretations example, a scenario using hypothetical maize
of the model, by either farmers or scien- varieties was created to better understand
tists, which can be sources of learning for the G×E interaction most valued by maize
both scientists and farmers (Cleveland and farmers in a study in Mexico, Cuba and
FIGURE 14.2
Ranking exercise as presented to farmers
Four types of maize are available
:
our farmer Modern variety (MV) our farmer variety with Modern variety with properties
variety (FV) properties from other from other organisms via
organisms via transgenesis transgenesis (TGMV)
(TGFV)
Rank these from best to worst for:
$ Planting in your fields
$ ou and your family to eat
Guatemala (Soleri et al., 2005). When asked and those same varieties as backgrounds for
to choose between two maize varieties a transgene: a transgenic farmers’ variety
with qualitative G×E response to annual and a transgenic modern variety (Figure
precipitation, 61 percent of farmers preferred 14.2). We asked farmers to rank these first
a variety with lower yield potential, mean as maize seed for sowing in their own fields,
and yield variation (‘stable’) to a variety and then again as maize grain for their
with higher yield potential, mean and yield family to eat. The FV and MV represented
variation (‘responsive’). The answer varied, two seed systems (informal vs formal,
with farmers from more difficult growing respectively) and had different agronomic,
environments preferring the stable variety at storage and culinary characteristics with
a significantly greater frequency than those which farmers were already familiar.
in more favourable growing environments. Farmers had no previous experience with
A similar scenario was created to investigate transgenic crop varieties (TGVs). Providing
farmers’ attitudes towards some of the these four choices allowed us to distinguish
possible consequences of pesticidal farmers’ preferences for varieties or genetic
transgenes in their maize varieties and the backgrounds (FV vs MV) from their
evolution of resistance in the pests that it preference for a genetic technology (TGV
controlled. Some of these consequences were vs non-TGV), an important distinction that
reliance on the formal seed system, a higher is either overlooked or confounded in most
seed price and initially high but declining research with farmers. TGVs were described
yields over time as pest populations evolved neutrally to farmers and they were given a
resistance. The hypothetical transgenic positive example of TGVs with the potential
variety was not identified as being transgenic to decrease pest damage.
when the scenario was presented to farmers.
Of those interviewed (n = 334), 70 percent 14.2 THE CONTEXT: INDUSTRIAL AND
chose a lower yielding but more stable and THIRD WORLD AGRICULTURE
locally available variety (Soleri et al., 2005). Industrial and Third World agriculture are
Similarly, an exercise asked those farmers to different in important ways in terms of seed
rank four types of maize: their own FV, a and food systems, growing environments
conventional MV they were familiar with, and crop genotypes. They are also similar in
FIGURE 14.3
Components of agricultural systems in traditionally-based
small-scale and industrial large-scale agriculture
Conservation ( in situ in fields Improvement (via propagule

and storage containers) selection for next year’s crop)
Consumption (primarily Production (primarily Multiplication (as part of

of food produced by for household crop production)
household) consumption)
a. Traditionally-based agricultural system: functions integrated in households and

communities
Improvement
Conservation
(by formal plant
(by scientists ex
breeding)
situ in gene
banks)
Consumption Multiplication
(by consumers (by seed institutes
distant from or companies)
production)
Production
(by specialized,
large-scale
farms)
b. Industrial agricultural systems: functions separated, specialized, many institutionalized

terms of the basic principles and processes holds sell some portion of their production
governing these variables and the interactions in the market, but they are incompletely
among them, including the outcome of integrated into these markets (Ellis, 1993).
choice and selection. Better understanding Farmers’ production knowledge combines
of these differences and similarities, and understanding based on theory and empiri-
their relationship to differences and cal observation with values about the social
similarities between farmers’ and breeders’ and cultural significance of farming, often
goals, knowledge and practices, can help to focused on FVs (Soleri et al., 2002).
further collaboration between farmers and Off-farm income is often critical for
plant breeders. households’ overall survival strategy, and
may reduce the importance of on-farm
14.2.1 Seed and food systems production. Migration of household mem-
In industrial agriculture, food production, bers, for example, may lead to labour short-
food consumption, crop improvement, seed age (Narayanan and Gulati, 2002) and to
multiplication and crop genetic resources reduced time and other resources devoted
conservation are specialized, physically to seed selection, conservation of crop
and structurally separated, and farming is genetic diversity or production, and even-
often considered to be primarily a busi- tually to loss of the knowledge on which
ness (Lyson, 2002) (Figure 14.3a). In SSTW they depend (for an example in central
agriculture these functions are combined Mexico, see Fitting, 2006).
within the farm household and community
(Figure 14.3b) (Soleri and Cleveland, 2004), Consumption
as described below. The differences due to Farm families rely on their own food
separation vs integration of these critical production for a significant proportion of
functions in seed and food systems have their food, and FVs are valued for traits that
important implications for decisions about contribute to storage, food preparation,
the best ways for farmers and breeders to taste, colour, texture and specific uses (e.g.
improve yields and quality traits, and to maize varieties grown for husks used in
minimize farmers’ risk. tamale production) (Soleri and Cleveland,
2001), or sticky rice FVs used for traditional
Production foods in southern China (Zhu et al., 2003).
SSTW agriculture is essential for feeding a These specialized uses mean some FVs have
significant proportion of the world popula- high market values.
tion now, and will probably remain so in
the future, even with production increases Improvement
in large-scale, industrial agriculture (Hazell Cultivation in new locations, farmers’
et al., 2007; Heisey and Edmeades, 1999). changing selection criteria and growing
As mentioned earlier, over 2 billion people environments were responsible for the
live on almost half a billion small-scale tremendous increase in intraspecific crop
farms (<2 ha) in the Third World, including diversity via mass selection following
half of the world’s undernourished people domestication (Harlan, 1992; Matsuoka et
and the majority of those living in absolute al., 2002) (see Chapter 1, this volume), and
poverty (Nagayets, 2005). Food production all of these continue today. It has been best
relies on household labour, and most house- documented at a local level in vegetatively
propagated crops (Elias et al., 2001), but using or selecting and saving FV seed each
also in predominantly self-pollinated crops year for planting (Louette and Smale, 2000;
such as rice (Dennis, 1987; Richards, 1986). Soleri, Smith and Cleveland, 2000). This
For cross-pollinating crops like maize, conservation is dynamic in that populations
farmers may not be interested in chang- are exposed to changing natural and artificial
ing quantitative phenotypic traits of their selection pressures, often creating locally
varieties through selection, but rather in distinct and adapted populations through
maintaining qualitative traits of interest indirect selection.
(Pressoir and Berthaud, 2004b), and can do Because food production and consump-
so successfully even in the presence of high tion and crop improvement, seed multi-
rates of gene flow at other loci (Louette, plication and conservation are all carried
Charrier and Berthaud, 1997; Pressoir and out within the same crop population, that
Berthaud, 2004a). Quantitative improve- population will not be optimized for any
ment in such species may more often be one function per se as it might be in indus-
sought through choosing new varieties or trialized systems. For example, the value of
populations, as discussed below. FV genetic diversity in its local conserva-
tion role may in some way be in conflict
Seed multiplication with the genetic composition ‘optimal’ for
Farmers do not usually distinguish seed its role as an improved population (Soleri
multiplication from food production, and Smith, 1995). In this sense, farmers’
although sometimes they plant separate crop populations are similar to semi-natu-
seed multiplication plots, as do some ral plant populations that
rice farmers in Sierra Leone (Richards, ...approach complex equilibria in
1986: 138–144). Farmers often save a high which overall fitness is as high as the
proportion of their seed from their own varied demands of differing sites and
harvests, but often also obtain seed through seasons, complex genetic control and
informal seed systems (Ndjeunga, 2002), the long term demands of adaptability
and frequently experiment with new seed allow.
(Louette, Charrier and Berthaud, 1997), (Simmonds and Smartt, 1999: 91)
including planting seed obtained as grain For this reason, and because of the value
(Soleri et al., 2005). The result is extensive of this farming for production and dynamic
gene flow via seed and other propagules, conservation, both in situ conservation by
as well as by pollen flow, creating seed farmers and ex situ conservation in formal
systems that are predominantly local and gene banks are necessary and complemen-
genetically open (Berthaud et al., 2001; tary. However, for conservation to play a
Pressoir and Berthaud, 2004a; vom Brocke useful role, interaction between farmers
et al., 2003a). and scientists is required, e.g. to ensure
that the selection environments in ex situ
Conservation conservation do not result in evolution that
Farmers conserve crop genetic diversity makes the population unsuitable for farm-
of FVs in situ in their fields and storage ers in the event that they require renewal
containers (Qualset et al., 1997). Most of their seed from outside their communi-
in situ conservation is done indirectly— ties (Soleri and Smith, 1995). In a similar
perhaps unintentionally—as a result of way, collaboration between farmers and
plant breeders needs to balance the goals with farmers, this environmental stress and
of the breeder, which will tend to focus on variation mean that selection for improved
improving specific traits, with the other performance in farmers’ environments
functions of the food and seed system. needs to take place in those environments,
and requires re-thinking some of the
14.2.2 Growing environments and assumptions of conventional plant breeding
genotypes (Ceccarelli and Grando, 2002). However,
Growing environments and crop genotypes many plant breeders, especially those with
of Third World farmers differ in impor- little experience with farmers’ growing
tant ways from those with which most environments, believe that as a general
plant breeders and agronomists in industrial principle selection should be done in
countries are familiar. Farms often consist optimal environments because there are
of a number of small, scattered fields with ‘spillover’ effects to marginal environments
marginal growing environments, i.e. rela- (for discussion, see Atlin et al., 2000;
tively high levels of stress and of temporal Rajaram and Ceccarelli, 1998). Thus,
and spatial variability. For example, while while plant breeders agree on the basic
the average size of maize grain farms in the principles of selection, they can disagree
United States of America in 2003 was 79.2 vehemently about how those principals
ha (USDA NASS, 2004), in the southern should be applied to farmers’ environments
Mexican state of Oaxaca over 76 percent of (Ceccarelli and Grando, 2002; Cleveland,
maize farms were smaller than 5 ha in 1995 2001). One source of such disagreements
(INEGI, 2001), and in one of the communi- may be differing interpretations of empirical
ties in the Central Valleys of Oaxaca where observations and theories thought to
we have worked, the average farm size is 3.7 underlie them (see Section 14.2.3).
ha and the average maize field size is 0.8 ha Farmers often continue to use locally
(Soleri, 1999; Soleri, Cleveland and Aragón selected FVs, even when MVs produced
Cuevas, 2003). In that same Oaxacan com- by the formal plant improvement and
munity, coefficients of variation of maize seed multiplication systems are available,
yields calculated using triangulation of because FVs may be better adapted to
farmer estimates were very high, averaging marginal growing environments, and
44 percent (Soleri et al., n.d.). Indeed it has because MVs may be agronomically,
been estimated that maize farmers in that culinarily and economically inappropriate
area of Oaxaca experience production fail- (Ceccarelli et al., 1994; Evans, 1993;
ure one year in four due to drought (Dilley, Heisey and Edmeades, 1999). Farmers
1997). In addition to high levels of environ- value FVs for agronomic traits, such as
mental variability, other factors contribute drought resistance, pest resistance and
to high levels of yield variability and pro- photoperiod sensitivity, as well as for traits
duction risk. SSTW farmers typically use contributing to storage, food preparation,
low levels of external inputs, and have lim- taste, market value and appearance (Smale,
ited access to government programmes and 2002). FVs include landraces, traditional
markets, and limited influence on the poli- varieties selected by farmers, MVs adapted
cies affecting them (Ellis, 1993; Hardaker, to farmers’ environments by farmer and
Huirne and Anderson, 1997). natural selection, and progeny from crosses
For many plant breeders who work between landraces and MVs (sometimes
referred to as ‘creolized’ or ‘degenerated’ the range of target environments to which

MVs) (Zeven, 1998; FAO, 1996). a genotype is adapted (Atlin et al., 2000;
FV yields are often much lower in the Bänziger and de Meyer, 2002; Ceccarelli
Third World compared with MV yields and Grando, 2002).
in industrialized agriculture, e.g. maize In specific situations, understanding this
yields in United States of America (∼8 t/ha) basic theory is difficult because a great
compared with Mexico (∼2 t/ha) (Aquino number of variables affect it, and predic-
et al., 2001) and Oaxaca (∼0.8–1.5 t/ha) tions are hampered by the lack of experi-
(Aragón-Cuevas et al., 2006). However, mental data and lack of the technologies
yield stability is often greater with FVs than and resources necessary to gather and ana-
for MVs grown in the same environments, lyse them. Plant breeders recognize that
because MVs often have steep response their theoretical understanding of plants
regression curves, i.e. are highly responsive beyond the basics is limited, and that much
to marginal environments, as well as optimal plant breeding has been based on intuition
ones (Ceccarelli, 1997; Evans, 1993). and empiricism rather than theory (Duvick,
An important reason for the higher yield 1996; Simmonds and Smartt, 1999; Wallace
stability of FVs is their higher level of genet- and Yan, 1998), although intuition and
ic diversity compared with most MVs, pre- empiricism are likely to be underlain to a
sumed to support broad resistance to multi- lesser or greater extent by the basic theo-
ple biotic and abiotic stresses (Brown, 1999). retical understanding of genotype x envi-
In addition, many centres of origin and cen- ronment relations.
tres of diversity for crop species are in the This fundamental biological theory is
Third World and cultivated primarily by the same no matter where plant breeding is
small-scale farmers, thus SSTW agriculture practised. However, the biophysical, eco-
is an important reservoir of genetic diversity nomic and sociocultural variables through
in the form of FVs (FAO, 1996). This diver- which this and other theories work can be
sity makes FVs valuable not only for farm- quite different. For example, think of the
ers, because they decrease the production contrast between farmers’ fields in marginal
risks in marginal environments, but also for environments and plant breeders’ research
the in situ conservation of crop diversity as a stations, or between national agricultural
source of resources for breeding MVs. policy priorities of large-scale efficiencies
and increased inputs and production, and
14.2.3 Plant breeder knowledge farmers’ priorities of reducing risk and
As outlined above (Section 14.1.3) the basic optimizing crop production as part of a
model of plant genotype-environment general household survival strategy. Work
interactions are well established and univer- under a specific set of circumstances may
sally accepted by plant breeders. However, lead to interpretation of theory that is then
many complexities of that model are still generalized and broadly applied, without
not well understood in terms of biological investigating the validity of those interpre-
theory (Duvick, 2002), and there continue tations under all circumstances. For exam-
to be disagreements about the interpreta- ple, the fundamental principle that—all else
tion of the basic model and its implications remaining constant—as E2 decreases, h2
for practice among plant breeders, such increases has been interpreted to imply that
as the effect of selection environment on selection in low E2 environments provides
the best response for all environments, Therefore, especially for those biophysi-
including ones with high E2 . However, cal aspects of genotypes and environments
empirical testing has shown this not to be that are less well understood in terms of
true in many cases (e.g. Ceccarelli, 1996; plant breeding theory, PBK may more
Ceccarelli et al., 1994, 2003; Comadran et likely to be based on each person’s or insti-
al., 2008); two reasons are that, first, the tution’s specific experiences with the par-
genes responsible for a quantitative trait ticular environments and crop genotypes
such as yield may be different in different they work with, and thus may be less gen-
environments (e.g. Atlin and Frey, 1990; eralizable, and more apt to be influenced by
Atlin, McRae and Lu, 2000; Venuprasad, pre-existing knowledge (including values)
Lafitte and Atlin, 2007), and, second, h2 of specific to the plant breeder’s social envi-
some important traits may not be entirely ronment. This means that disagreements
obscured by E2 (Al-Yassin et al., 2005) between farmers and plant breeders, and
(see Chapter 2, this volume). Working among plant breeders, could arise even
with farmers often requires that breeders though fundamental genetic and statistical
test the validity of those interpretations of principles remain constant across a range of
theory that form the basis of conventional contexts, because the ‘art’ of plant breeding
plant breeding. This includes comparing is more tied to specific individuals or envi-
the genotypes and environments and goals ronments (Ceccarelli and Grando, 2002;
for improvement, and testing the assump- Soleri and Cleveland, 2001).
tions (biological, environmental, economic,
sociocultural) on which they are based, 14.2.4 Farmer knowledge
and adjusting interpretations of theory, A lack of empirical research and theoretical
and hence methods used (Ceccarelli and analysis has contributed to using overly sim-
Grando, 2002). plified definitions of FK (and often of PBK
For this reason, farmer–breeder col- as well), and the common failure to test the
laboration may often benefit from making many assumptions underlying these defini-
a clear distinction between (a) fundamental tions (Cleveland, 2006; Sillitoe, 1998). We
biological theory, (b) interpretations of can very roughly divide current views of FK
fundamental theory, and (c) methods and into two categories: there are those that see
practice, with ‘c’ possibly very different FK as fundamentally different from PBK,
depending on whether it is based on ‘a’ or and those that see it as fundamentally similar.
‘b’, or on different versions of ‘b’. Many These views also form the basis of particu-
of the disagreements about plant breeding lar advocacy perspectives; generally neither
methods for participatory plant breeding considers the theoretical content of FK.
(PPB) may grow out of disagreements In the first category, definitions of FK
about differences in the interpretation of emphasize that it is primarily value-based,
fundamental biological theory, and disa- comprising intuition and skill, socially con-
greements about these interpretations may structed, and based on the local social
in turn be based on the belief of proponents and environmental contexts and culture.
that their interpretations of fundamental According to this perspective, farmer and
theory are not based on their unique expe- PBK are seen as fundamentally different,
riences and assumptions, but rather are part and attempts to explain FK in scientific
of fundamental theory. terms impede true appreciation of FK.
The second category emphasizes appears to have been “better understanding

that FK consists primarily of rational of new ideotypes based on farmers’ expe-
empirical knowledge, usually focusing on riences, specific preferences and needs”
either economic or ecological knowledge. that will affect priorities of formal plant
Definitions of FK as economically rational breeding and the “process of formal variety
tend to assume that scientists are more development” (Weltzien et al., 2003: 75).
rational, and that farmers are risk neutral More recently, using FK of crops as
and their behaviour is based on a desire for a discriminatory tool has become more
profit maximization in the form of high common. This has been important in some
average yields (e.g. Zilberman, Ameden and PPB work, with farmers asked to choose
Qaim, 2007). According to this definition, among varieties already released in other
the role of outsiders should be to facilitate areas (e.g. for rice and chickpea; Joshi
the replacement or modernization of small- and Witcombe, 1996), among new and
scale farming, including replacement of FVs experimental varieties (e.g. for pearl millet;
with MVs (Mohapatra, Rozelle and Huang, Weltzien et al., 1998), or among segregating
2006; Srivastava and Jaffee, 1993). The populations (e.g. F3 bulks with barley;
definition of FK as ecologically rational Ceccarelli et al., 2000), or to select individual
tends to assume that farmers have detailed, plants within segregating populations (e.g. F5
accurate and therefore sustainable ecological bulks of rice; Sthapit, Joshi and Witcombe,
knowledge of their environments. The first 1996; and F4 bulks of rice; Virk et al.,
part of that definition is supported by much 2003). When such choice or selection is
empirical data, especially ethnotaxonomic accomplished using actual plants, plant parts
studies of plants and animals, while or propagules, analysis of results can reveal
recognizing variation in distribution of farmers’ implicit criteria that they may not
cultural knowledge as the result of factors be able to verbalize easily, if at all (i.e. it may
including age, gender, social status and be unconscious) (Louette and Smale, 2000;
affiliation, kinship, personal experience and Soleri, Smith and Cleveland, 2000).
intelligence (Berlin, 1992). These approaches to understanding FK
Participatory research has usually been have made valuable contributions to achiev-
based on the second definition of FK. As a ing more effective crop improvement for
result, the focus in using farmer knowledge farmers’ conditions. However, the theo-
has been on the details it can provide in the retical basis of FK is not usually considered,
form of a discriminatory or, most frequent- and rigorous comparisons with PBK have
ly, descriptive tool in PPB. For example, a not been carried out, “opportunities rarely
major survey of 49 PPB projects found that develop for interaction between breeders
the primary focus was soliciting farmers’ and farmers beyond the survey”, with the
descriptions and rankings of selection crite- discussion “driven by the breeders’ con-
ria. For about two-thirds of these projects, cepts of the present situation, making it
“identifying, verifying, and testing of spe- difficult for farmers to express their views
cific selection criteria was the main aim in the context of their reality” (Weltzien
of the research”, and 85 percent obtained et al., 2003: 51). It may also be difficult
farmers’ selection criteria for new varieties for farmers to communicate to outsiders
(Weltzien et al., 2003: 17–18, 51, 75). The their knowledge that goes beyond descrip-
main impact on scientific plant breeding tion or discrimination. For this reason a
definition of knowledge—both farmer and farmers continued as before. There have

scientist—as complex, and including values, also been marked changes in crop genetic
empiricism, theory and experience is useful diversity over time, especially at specific
(Cleveland, 2006). This definition underlies and intraspecific taxonomic levels.
an approach that starts with basic theoretical
knowledge and clearly distinguishes theory 14.3.1 Domestication and subsequent
from its local interpretation, in an attempt changes in diversity
to better understand farmers’ choice and While domestication resulted in a large
selection, and to identify possible bases for decrease in the number of plant species
substantive collaboration between farmers exploited, it was followed by large increases
and scientists. In the rest of this chapter in intraspecific diversity, as FVs evolved as a
we use this definition to look at two key result of natural and artificial selection in new
processes in plant breeding: choice of popu- biophysical and sociocultural environments
lations (or varieties) for direct use or for (Harlan, 1992) (see Chapter 1, this volume).
further breeding, and selection of individu- For many of the more widely grown food
als within a population. We focus on our crops, domestication resulted in evolutionary
understanding of FK and practice of choice changes making them genetically distinct
and selection, how farmers and scientists from their closest wild relatives today, and
can better collaborate in those steps, and most became dependent on humans for
why such collaboration is important. reproduction (Harlan, 1992; Simmonds and
Smartt, 1999). Exceptions exist, especially
14.3 FARMER CHOICE AND SELECTION: among some perennial fruit crops, more
PAST, PRESENT AND FUTURE accurately described as semi-domesticates,
While this chapter is primarily about con- where crops are not the result of selection
temporary farmer and plant breeder choice resulting in E, but rather are choices of
and selection, a brief look at the broad superior genotypes from among those extant
trends in the past, present and future of crop in the wild (for olive, see Baldoni et al., 2006,
improvement in relation to farmers will and Breton et al., 2006).
help in understanding the challenges and Domestication seems likely to have
potential for plant breeding with farmers. been the result of indirect selection and
This section is not essential for understand- unintentional direct selection (e.g. when
ing the rest of the chapter, and might be farmers select for large seed size or brittle
quickly skimmed and used as a reference. rachis as a result of their seed collection
As measured by the rate of desired behaviour; Harlan, 1992), and perhaps
crop genetic changes achieved by selec- some intentional selection for evolutionary
tion, three broad stages have been sug- change (see Section 14.1.2). However, it is
gested (Gepts, 2004). Initial rapid progress very difficult or impossible to determine
with domestication was followed by long the type of selection that resulted in past
periods of much slower change as original crop evolution, and experts differ on the
domesticates spread to new environments type they believe was most important.
and responded to a range of new natural For example, Allard emphasizes direct,
and artificial selection pressures, and with intentional selection,
modern plant breeding the rate of change The consensus is that even the earliest
in MVs increased substantially, while most farmers were competent biologists
who carefully selected as parents tion) and other linked loci. The presence
those individuals … with the ability of this sweep distinguishes the sticky rice
to live and reproduce in the local favoured by upland northeast Asian peo-
environment, as well as with superior ples from the non-glutinous rice varieties
usefulness to local consumers. used by other Asian groups, and presum-
(Allard, 1999) ably would be among their fundamental
In contrast, Simmonds and Smartt (1999: 13) choice criteria, perhaps as an adaptation for
emphasize indirect selection: “the art of eating with chopsticks (Olsen et al., 2006).
cultivation is perhaps the peasant’s most Increasing evidence for a number
potent contribution.” of crops suggests that domestication
Similar to studies based on archaeologi- could have occurred over short periods
cal data, results of molecular analyses sup- relative to the ~12 000 years that crop
port the hypothesis that farmers’ selection plants have been cultivated (Gepts, 2004).
has been successful in achieving evolution- Domestication syndrome traits often
ary change for traits in the ‘domestication appear to be determined by a small number
syndrome’ that might be indirectly or unin- of genes with large effects, suggesting that
tentionally favoured because of agronomic domestication could proceed relatively
superiority (see Chapter 1, this volume). rapidly. For example, Paterson et al. (1995)
There is also evidence that farmer selection found a small number of quantitative trait
has been a powerful force for evolutionary loci (QTLs) coding for the domestication
change based on other preferences as well. syndrome traits of seed size, photoperiod
For example, three major genes involved in sensitivity of flowering, and brittle rachis
starch metabolism in maize were found to in taxonomically distinct cereals with
have unusually low genetic diversity com- diverse centres of origin (sorghum, rice
pared with its closest wild relative (teos- and maize). In common bean (Phaseolus
inte, Zea mays subsp. parviglumis), which vulgaris L.), control of the domestication
is strong evidence of selection for specific syndrome involves genes that have a large
processing and culinary qualities impor- effect (>25–30 percent) and account for a
tant for the primary manner in which substantial part of the phenotypic variation
maize has been consumed in its regions of observed (>40–50 percent) (Koinange,
origin and diversity (Whitt et al., 2002). In Singh and Gepts, 1996). Simulations based
addition, three other loci contributing to on sequence variations at loci coding for
sweet maize grain phenotypes showed low biochemical or structural phenotypes in
diversity (resulting from strong selection) maize and its close and distant relatives
in only certain varieties in particular loca- have estimated that domestication
tions, evidence of further specialization could have taken from 10 (Eyre-Walker
in the non-agronomic selection pressures et al., 1998) to between 315 and 1 023
farmers have exerted on maize (Olsen et generations (Wang et al., 1999). In
al., 2006; Whitt et al., 2002). Similarly, it addition to selecting for characteristics of
appears that strong directional selection the ‘domestication syndrome’, especially
for sticky, glutinous grain quality resulted in cereals and small pulses (Harlan, 1992),
in a selective sweep affecting an area over domestication in sexually propagated crops
250 kb long that includes the locus cod- may have resulted in increased autogamy
ing for this quality (low amylase produc- and therefore homozygosity, expressed
phenotypically in greater trueness to farmers working in combination with

type in a population over generations. local natural selection contributed to the
In contrast, some vegetative propagation large amount of intraspecific diversity that
may have selected for heterozygosity (via evolved following domestication:
heterosis) and therefore for allogamy, Probably, the total genetic change
as contemporary evidence suggests for achieved by farmers over the millennia
cassava (Pujol, David and McKey, 2005). was far greater than that achieved by
The genetic changes that define crop the last hundred or two years of more
domestication are inextricably linked systematic science-based effort.
with changes in selection pressure. (Simmonds and Smartt, 1999: 12).
These pressures are not only exerted by
direct human selection of propagules for 14.3.2 Modern, scientific plant
planting, but perhaps more often with the breeding
differences in selection pressures created Farmer and plant breeder crop improvement
by human modification of growing began to be separated about 200 years ago in
environments (Figure 14.1). In southeast “technically advanced temperate countries”
China, for example, evidence for the earliest (Simmonds and Smartt, 1999: 12) with the
cultivation of both wild and domestic rice beginning of specialized, amateur breeding.
(~7 700 BPE) suggests that this occurred The widespread acceptance of evolution and
where farmers were intensively managing the rediscovery of Mendel’s research after
coastal wetlands with fire to control 1900 eventually led to modern scientific
vegetation and bunds to control flooding, plant breeding, based on a combination
and increased nutrient concentration in of Darwinian evolution, Mendelian
fields (Zong et al., 2007). Bringing wild genetics and biometry (Fitzgerald, 1990;
plants into human modified environments, Provine, 1971), with modern plant
such as compost heaps near houses, as breeders considering themselves ‘applied
well as exchange of seeds and other evolutionists’, whose goal is to develop
propagules, also facilitated domestication plant varieties better adapted to growing
via hybridization, as with Leucaena in environments, measured primarily as
southern Mexico, and probably with two increased yield (Allard, 1999).
other important domesticates from that Farmers and formal plant breeders con-
region, agave (Agave spp.) and prickly-pear tinued to collaborate at this time, for exam-
cactus (Opuntia spp.) (Hughes et al., 2007). ple in making crosses and selections in maize
Domestication generally decreased the breeding in the United States of America
fitness of plants in natural environments, (Fitzgerald, 1990; Schneider, 2002). But as
and made them more dependent on humans the importance of evolutionary theory in
and human-managed environments. plant breeding increased in comparison
The geographical spread of domesticated with empirical heuristics, the economic
crops led to great varietal diversification importance of plant breeding increased and
as a result of the increase in diversity of came to dominate formal plant breeding by
natural and artificial selection pressures professional plant breeders. Simultaneously,
encountered, followed by choice among the farmer’s role in crop improvement in
preferred populations. It is generally industrial countries decreased, for example
assumed that simple mass selection by in the United States of America (Fitzgerald,
1990; Kloppenburg, 1988) and Switzerland industrial agriculture, and is expanding in

(Schneider, 2002). Plant breeders’ concepts the Third World. Of the 23 countries grow-
subsequently developed independently of ing TGVs in 2007, 12 were ‘developing’
farmers’ concepts, effectively separating countries, and estimated to account for 43
the formal from the informal systems of percent of the area planted and 90 percent
crop improvement and seed multiplica- (11 million) of the farmers growing TGVs.
tion. When farmers are involved by con- Of these, 99 percent (10.9 million) were in
temporary plant breeders in their work China and India, growing mostly Bt cot-
it has generally been limited to the stage ton (James, 2006, 2007). Currently, TGVs
of evaluating and choosing among plant of food crops for Third World farmers are
breeders’ populations or varieties in their either planned, being developed, in field tri-
fields (Duvick, 2002). als, or approved and in production.
TGVs are currently being promoted by
14.3.3 Biotechnology development organizations, governments
Advances in genetics and molecular biol- and corporations as the key to increasing
ogy have led to developments in biotech- production and income and reducing hunger
nology that have dramatically enhanced and malnutrition in SSTW agriculture
the ability to understand and manipulate (FAO, 2004; Rockefeller Foundation,
plant genomes. Functional genomics has 2007; World Bank, 2007). However, the
elucidated the relationship among genetic focus on TGVs to improve Third World
components and to phenotypes; marker agriculture is very controversial (Abate
assisted selection (MAS) has increased the et al., 2008; Stokstad, 2008). A number of
efficiency of breeding for specific traits; and studies, mostly by economists and of Bt
genetic engineering has made it possible cotton, maize and rice, have concluded
to transfer genes from almost any organ- that farmers readily adopt TGVs because
ism into a crop species. When these genes they increase yield and income, reduce
come from a different species the process pesticide applications or improve farmer
of transformation is called transgenesis, health (Gouse et al., 2006; Huang et al.,
and the resulting crop variety a genetically 2003, 2005; Morse, Bennett and Ismael,
engineered (GE) variety, genetically modi- 2006; Qaim and Zilberman, 2003). Other
fied organism (GMO) or, most accurately, a studies have found that adoption may be
transgenic crop variety (TGV). the result of fads (Stone, 2007) or a lack of
TGVs are a rapidly growing agricul- freedom to choose (Witt, Patel and Schnurr,
tural technology, with the area planted 2006), and that higher yields and reduced
increasing by 9.4 percent from 2007 to pesticides may be reversed after several
2008, to over 125 million hectares (James, years due to the emergence of secondary
2006, 2008), or over 9 percent of cultivated pests (Wang, Just and Pinstrup-Andersen,
land globally (calculated from FAO, 2007, 2006). Others have suggested that the net
2009). Currently grown TGVs are prima- benefits of TGVs may not be as great
rily targeted to industrial agriculture and as those of alternative improvements in
designed to enhance yield and net profit for agriculture (e.g. Uphoff, 2007). The potential
farmers by directly reducing pest damage ecological and genetic effects of TGVs and
or facilitating herbicide use. Globally, most transgene flow into non-TGV crop or wild
of the area planted to TGVs is in large-scale or weedy populations, especially in Third
World agriculture, are not well understood traditions or current needs (Cleveland and
(Ellstrand, 2003b; Heinemann, 2007; NRC, Murray, 1997).
2002; Snow et al., 2005). Much plant breeding has moved from
The spread of biotechnology has also the public to the private sector (Frey, 1996)
resulted in unintentional transgene flow, and thus selection criteria are increasingly
including into centres of diversity, e.g. maize vulnerable to being dominated by private
transgenes documented in Mexican FVs profit motives rather than public good
(Alvarez-Morales, 2002; Pineyro-Nelson et motives (Simmonds, 1990), which is espe-
al., 2009; Serratos-Hernández et al., 2007). cially evident for TGVs. The major share
Such transgene flow can be difficult to pre- of agricultural biotechnology processes and
vent (NRC, 2004), the early stages of trans- products are controlled by private multina-
gene flow to FVs are extremely difficult to tional corporations with little incentive to
monitor (Cleveland et al., 2005), and the develop TGVs most appropriate for Third
effects may often be irreversible (Ellstrand, World farmers who cannot afford to pay
2003a). Potential effects of transgene flow the premium for TGV seed (CGIAR, 2006;
on FVs and farmers are both positive and World Bank, 2007: 178).
negative, and will require risk analysis and Similarly, there is increasing concentration
evaluation specifically adapted to each loca- in the seed sector, which potentially
tion – crop combination within the Third reduces competition and limits the kinds
World (Cleveland and Soleri, 2005; Soleri, of crops and crop varieties produced and
Cleveland and Aragón Cuevas, 2006). made available. The largest seed companies
Transgenes can introduce novel forms of control an ever larger proportion of the
diversity into the crop populations being seed market; according to one estimate,
selected upon by farmers and plant breeders, between 1997 and 2004 the companies with
but there is no reason to expect that farmers the largest sales increased their market share
will be able to retain, discard or manipulate from 27 percent to 33 percent, and in 2004
them any differently from other genes. the top four companies owned 38 percent
of biotechnology patents (World Bank,
14.3.4 Privatization 2007: 135–136).
In the early 1980s, some countries and The drive to globalize industrial-world
farmer support groups sought to do away IPRs in plants has been intensified as a result
with all intellectual property rights (IPRs) of pressure from agricultural biotechnology
in crops, establishing ‘farmers’ rights’ to all corporations (Graff et al., 2003; Shorett,
crop genetic resources, but this move was Rabinow and Billings, 2003). This means
defeated by the United States of America that as patented TGV crops and their trans-
and other industrial nations (Fowler, 1994), genes move intentionally or unintentionally
and private rights in plants and other living around the world, so could the rights of the
organisms now dominate, with industrial companies who own them. Movement of
patents leading the way (Atkinson et al., transgenes into non-transgenic crop popu-
2003). Farmers were left with having to lations, whether producing a net benefit for
defend themselves from the advances of an the farmer or not, makes farmers vulnerable
IPR system in plants designed by industrial to IPR claims from the technology devel-
nations and corporations, a system that oper. In the United States of America and
generally does not recognize farmers’ many other industrialized countries, patent
holders have rights to seek damages from 14.3.5 Sustainability and farmer-
farmers who end up with patented genes scientist collaboration
in their crops, even though farmers do not The search for sustainability provoked by
want them, and do not know they are there negative environmental impacts of agri-
(Janis and Kesan, 2002). The World Trade culture (Matson et al., 1997; NRC, 2002;
Organization (WTO) seeks worldwide Tilman et al., 2002), and its genetic vulner-
uniformity of laws for IPRs in plants and ability (NRC, 1991) has led to the incor-
plant DNA to facilitate global enforce- poration of more genetic variation within
ment, and many Third World countries and among varieties by the formal crop
have adopted the industrial world model improvement system (Cooper, Spillane and
(UPOV – International Union for the Hodgkin, 2001; Ortiz et al., 2007). It has
Protection of New Varieties of Plants) also encouraged a re-evaluation of G×E
while others have adapted their national in crops and how best to exploit this
laws to protect small-scale farmers (World for farmers’ conditions (Ceccarelli et al.,
Bank, 2007: 167). The spread of IPRs and 1994, 2001). The impact on farmer selec-
coupled economic control of agricultural tion will be in the greater intraspecific and
biotechnology means that Third World intravarietal diversity deployed in formally
farmers and the nation states they live in developed varieties, and greater interest in
will have a difficult time gaining mean- that system for breeding goals more simi-
ingful control of the means to intention- lar to those of farmers. Part of the interest
ally create TGVs more suited to their own in sustainability (environmental, economic
needs, if this is the path they choose. As and social) has led to collaboration between
a result, most organizations promoting farmers and scientists.
TGVs more suited to Third World farmers Participatory or collaborative plant
are advocating public-private partnerships breeding is attempting to reverse the
(CGIAR, 2006; FAO, 2004; World Bank, separation of farmers and scientists and
2007), yet it is not clear how farmers’ improve the outcomes of choice and
rights will fare in this collaboration, and selection in farmers’ terms (Cleveland
they are not being rigorously addressed. and Soleri, 2002a; PRGA, 2004; McGuire,
While most corporations deny they Manicad and Sperling, 2003; Weltzien et
would enforce their IPRs against Third al., 2003). To that end, the next sections
World farmers, there are no guarantees. focus on understanding farmers’ choice
In addition to transgenes, control of local and selection, and thereby enabling farmers
farmers’ crop genetic resources, and the and scientists to work more closely and
traditional names and other cultural prop- productively in improving the crops they
erty that go with them through industrial grow and depend upon.
IPRs, can legally and economically pre-
vent local people themselves from reaping 14.4 CHOICE OF GERMPLASM
potential benefits in a global marketplace It is important for plant breeders to under-
increasingly interested in traditional crops stand how and why farmers choose varie-
and foods (Soleri et al., 1994). There are ties of their crops, because farmer choice
already cases of this, some of which are will ultimately determine whether a new or
being challenged (Pallotini et al., 2004). improved variety will be useful. In this sec-
tion we consider choice based on perceived
risk and yield stability, and on other fac- farmers’ fields, or even within a field (Soleri
tors, including quality traits. et al., 2002). Variation in time is also large:
Just as there are factors favouring the in the semi-arid tropics, seasonal and
inclusion of more than one variety in a annual rainfall is highly variable, and even
farmer’s crop repertoire, there also are fac- in years with adequate total rainfall, rains
tors limiting the number of varieties chosen. may arrive late, end too early, stop for a
These include farmers’ resources, growing period or be too heavy during flowering
environments and crop reproductive biology, or harvesting. Therefore farmers may often
among other possible factors. Additionally, grow two or more varieties of many crops,
if crop varietal diversity is maintained at a each with distinct agronomic characteristics
community instead of household level, then presumably “as a measure of insurance
farmers may not feel the need to maintain against vagaries of the weather, diseases, or
some varieties themselves each year, even pests” (Doggett, 1988).
though they consider those varieties to be Understanding farmers’ choice can
part of their varietal repertoire and intend to provide valuable insights for scientific plant
grow them in the near future (for the case of breeders. In response to climate change
rice, see Dennis, 1987). in the form of the southern movement of
isohyets, policy-makers in Mali argue that
14.4.1 Varietal choice, yield stability improved short-cycle varieties are a critical
and risk part of stabilizing the country’s volatile
In much of the past plant breeding for cereal production (Dembélé and Staatz,
SSTW farmers, it was assumed that high 2000). One result is that both sorghum
yielding varieties selected in more optimal breeders and farmers in southern and central
environments would outyield FVs in Mali look north for shorter cycle varieties.
farmers’ environments (Ceccarelli et al., Interviews in four villages in the Upper Niger
1994; Ceccarelli, Grando and Booth, 1996). River valley zone of Mali found the most
If farmers did not adopt these varieties it common reason for adoption of the three
was assumed that they were ignorant of most popular sorghum varieties was early
how to improve their growing environments maturity (Adesina, 1992). However, since
(Aquino, 1998), or if they could not afford in good rainfall years long-cycle varieties
to do so, it was assumed that they should generally have higher yields (Adesina, 1992)
get out of farming. Consideration of risk and are rated higher for quality (Ingram,
provides a different understanding of Roncoli and Kirshen, 2002), farmers do not
farmers’ varietal choices and other practices. give these up entirely. Their choices thus
In the conventional economic model, a risk- increase the number of varieties in their
neutral farmer would only grow the one repertoires, although the net impact on
variety that gives the highest profits per unit genetic diversity has not been investigated.
area (Smale, 2002). However, many small- Another study in Mali of farmers’ choices
scale farmers in marginal environments are among their traditional sorghum varieties
risk averse (Anderson and Dillon, 1992; Soleri in terms of one or more than one variety,
et al., n.d., 2008), and spatial environmental and short-cycle or long-cycle varieties,
variation increases the likelihood of cross- found that farmers make these choices in
overs in varietal performance (qualitative an effort to optimize outputs in the face of
G×E; see Section 14.5.1, below) between variation in the growing environment and in
availability of human-managed inputs, such mental variation is minimal, there may be

as labour and tools. For example, better little incentive for farmers to maintain FVs
rains in 2002 compared with 2001 appear to while adopting MVs in order to reduce risk
be a major factor in the general shift toward due to qualitative G×E (Virk and Witcombe,
a greater number and longer cycle length of 2007). Clearly, the diversity outcome of
varieties, with 60 percent of farmers adding locally focused improvement programmes
varieties between 2001 and 2002 (Lacy, will depend on the specific situation.
Cleveland and Soleri, 2006).
The need for research on farmer choice 14.4.2 Other factors influencing choice
and risk is also illustrated in the case of Farmers may also choose more than one vari-
potato in the Andes (Zimmerer, 2002). An ety because of their different quality traits.
emphasis on potato varieties with large For example, interviews with 599 Nigerian
tubers because farmers prefer the higher farmers supported the conclusion that they
yield of these varieties would ignore the grow both long-cycle and short-cycle cowpea
fact that poorer farmers actually select varieties: short-cycle for food grain and long-
small tubers for planting because they cycle for feed during the dry season when
can reduce the amount of potential food other fodder sources are scarce (Abdullahi
used for planting material. An implication and CGIAR, 2003). Some maize farmers in
is that the varieties poor farmers would Oaxaca, Mexico, maintain varieties specifical-
actually choose to plant may be quite ly for their coloured husks or tassels because
different from that anticipated by breeders, of their aesthetic qualities, e.g. coloured husks
indicating changes were needed to make used to wrap tamales impart their colour to
improvement programmes more relevant them (Soleri, field notes, 1996–1999), and
for those farmers’ needs. families who make the traditional beverage
These and other studies suggest that tejate maintain more varieties of maize, using
crop improvement programmes need to spe- them in its preparation (Soleri, Cleveland and
cifically target farmers’ growing environ- Aragón Cuevas, 2008).
ments and needs, and use local germplasm The number of varieties grown by farm-
as the basis for this (Ceccarelli and Grando, ers may also be influenced by seed source
2002). They indicate the importance of and social variables (David, 2004). In a study
plant breeders supporting varietal portfo- of Mexican maize farmers, choice of total
lios (Ceccarelli et al., 2003; vom Brocke et number of varieties grown was related to
al., 2003a; Weltzien et al., 2003) available household seed source. Households plant-
through farmer-to-farmer exchange as an ing mostly their own seed chose an average
alternative to the development of a small of twice as many varieties in comparison
number of varieties for large-scale adoption. with those households that obtained all
In addition to decreasing farmer risk, this their seed from non-household sources
strategy also supports conservation of crop (Louette, Charrier and Berthaud, 1997). In
genetic diversity in situ (Ceccarelli, Grando a review of field research on farmer crop
and Baum, 2007). However, there is also genetic resources, wealth was a common
some evidence that MVs developed through indicator for producers who cultivated
participatory varietal selection can replace more varieties compared with resource-
existing FVs, as with wheat in South Asia poor producers (Jarvis et al., 2000). The
(Ortiz-Ferrara et al., 2007). When environ- choice of total number of sorghum varie-
ties may be significantly related to ethnic- of E2 and G2 to P2 , and if they distin-

ity, as in one area of the United Republic guished between high and low h2 traits in
of Tanzania, where migrant Gogo farmers their major crop. The first null hypothesis
from a traditional sorghum-growing region was that there was no difference in distri-
grow more than twice the number of varie- bution of farmers’ responses concerning
ties as migrant groups from maize-growing consistency between parent and progeny
regions (Friis-Hansen and Sthapit, 2000). phenotypes in a typical, variable environ-
ment and in a hypothetical, uniform envi-
14.5 SELECTION ronment for (i) relatively low h2 traits, and
Given the historical background outlined (ii) relatively high h2 traits. This hypothesis
earlier, including the emphasis on selection was rejected for low h2 traits, but accepted
as practised by scientists, we now discuss for high h2 traits (most farmers anticipated
the concept and process of selection, no change in phenotype regardless of envi-
emphasizing the contexts and perspectives ronment), suggesting farmers see little or
of farmers. We begin by reviewing research no contribution of genotype to P2 for low
on farmer understanding of heritability h2 traits, and the opposite for high h2 traits.
and G×E, two fundamental concepts in The second null hypothesis, that farmers’
selection. responses indicate no perception of differ-
ences in h2 for relatively low and high h2
14.5.1 Farmer understanding of trait expression in a variable environment,
heritability and G×E was also rejected, supporting the conclu-
Heritability (h2) is a key determinant of sion that farmers do perceive differences in
genetic response (R) (see Section 14.1.3). h2 of traits. Thus, most farmers distinguish
One of the main factors that decreases h2 between high and low h2 traits, and con-
is environmental variability ( E2 ). Another sciously select for the former, while often
important and related element affecting the considering it not worthwhile or even pos-
outcome of selection is G×E. Interpretation sible to seek R > 0 for the latter, especially
of G×E will influence plant breeders’ in cross-pollinated crops (Soleri et al., 2002).
approaches to developing and improving Given farmers’ experiences and the tools
crop varieties and their choices of how and methods available to them, the role
many and which varieties will be released of G2 in low heritability traits is obscured
across agricultural environments (Cooper by the E2 in their growing environments.
and Hammer, 1996). For these two impor- Similarly, Ceccarelli (1996) argues that plant
tant elements that affect the results of breeders’ lack of experience with growing
selection, experience as well as goals will environments as stressful and variable as
influence the knowledge of farmers and those of farmers has obscured plant breed-
plant breeders and how each responds to ers’ ability to perceive qualitative G×E in
variations in h2 and G×E in their crop vari- some MVs between farmers’ environments
eties and growing environments. and the more favourable ones they are
In comparative research on farmers’ con- accustomed to.
cepts of h2, farmers were presented with To understand farmers’ perceptions
scenarios about both high and low h2 traits of spatial G×E interactions we used a
(Figure 14.4, Table 14.1). The goal was to scenario with two genotypes originating in
determine if farmers noted the contribution contrasting growing environments at three
FIGURE 14.4
Sample scenario used to elicit farmers’ knowledge of heritability
Seed from long ears only is planted
Typical local field, variable and high stress Hypothetical uniform and optimal field
For each field we asked, “What length will the ears that grow in this field
be? The same as or different from the ears from which the seed was taken?”
This scenario, used with farmers in Cuba and Mexico, is about a maize trait with low average heritability (ear length). We
asked farmers what the ear length of the next generation would be in a typical relatively variable, stressful field and a
hypothetical uniform, optimal field if seed from long ears only was planted. Similar scenarios were used for low and high
heritability traits for crops in the five sites in study (see Table 14.1).
TABLE 14.1
Understanding farmers’ perceptions of heritability
Location, crop Null hypothesis #1: Null hypothesis #2:
For traits with relatively low or those with relatively In scenarios depicting a typical
high h2, distribution of farmers’ responses is the environment, distribution of farmers’
same whether scenarios depict typical or optimal responses is the same for traits with
environments, i.e. farmers do not see a contribution relatively low or those with relatively
of environment (Env) to phenotype high h2, i.e. farmers do not see a
difference in h2 between traits
(a) Low h2 trait across (b) High h2 trait across Low v high h2 traits in typical, variable
typical and optimal Envs typical and optimal Envs Env
Cuba, maize Ear length* Husk colour Ear length v. husk colour*
Mexico, maize Ear length* Tassel colour Ear length v. tassel colour*
Mali, sorghum Panicle weight* Glume colour* Panicle weight v. glume colour*
Syria, barley Plant height* Seed colour Plant height v. seed colour*
Nepal, rice Plant height* Seed colour Plant height v. seed colour*
* Hypothesis rejected, Fisher’s exact test, P<0.05
Based on Soleri et al., n.d.
levels: between locations, between fields evolution, breeders looked to farmers for
in one location, and between places in one their applied knowledge and practice that
field (Soleri et al., n.d., 2002). The results produced practical results in the form of
indicated that farmers (n = 208) perceive new varieties, as in the early commercial
inter- (57 percent) and intra- (30 percent) development of maize in the United States of
location G×E for their major crop, though America (Wallace and Brown, 1988: 87–90).
far fewer at the latter level. G×E within With the increased importance of formal
a field (18 percent) was noted mostly, science in plant breeding compared with
though not exclusively, by those growing empirical heuristics, and later as plant
self-pollinated crops, and especially those breeding moved from the public to the
working at a small scale with intimate private sector (Kloppenburg, 1988), plant
knowledge of within-field soil and breeders began to eliminate farmers from
moisture variations (e.g. rice farmers in their work (e.g. Schneider, 2002). Plant
western Nepal). Similarly, 37 percent of breeders’ and farmers’ practices and concepts
farmers responded that a qualitative G×E subsequently developed independently of
interaction could occur in their crop due each other, effectively separating the formal
to temporal environmental variation in and informal systems of crop improvement
the form of annual precipitation. In the and seed multiplication, with plant breeders
presence of qualitative G×E, perceptions coming to dominate: “a trend that has been
of the best genetic strategy may differ at least locally apparent for 200 years”
and be informative about the needs of (Simmonds and Smartt, 1999: 13). Plant
particular groups or regions. As mentioned breeders focused on modern varieties widely
in Section 14.4.1 above, farmers tended to adapted to more optimal, more intensively
favour yield stability over high yield when managed environments, while many
choosing among varieties in the face of traditionally-based farmers in relatively
qualitative temporal G×E. This choice was marginal environments continued to focus
significantly more frequent among farmers on traditional, specifically adapted varieties
in more difficult environments compared for their diverse, more marginal growing
with more favourable environments. environments (Ceccarelli and Grando, 2002;
Cleveland, 2001). When contemporary plant
14.5.2 Farmers’ selection goals breeders involve farmers in their work, it
If plant breeders misunderstand what has generally been limited to the stage of
farmers are and are not attempting to evaluating the plant breeders’ populations
accomplish with their selection practices, or varieties in the field (Duvick, 2002), i.e.
it can limit the potential for meaningful choosing among different populations or
collaboration and lead to inappropriate varieties, not selecting among different plants
investments of scarce time and resources. to genetically change existing populations
Such misunderstandings have grown out or varieties.
of the historical process of separation Today, many modern plant breeders
of farmers’ and plant breeders’ work consider themselves to be ‘applied
(Cleveland and Soleri, 2007). evolutionists’, whose goal is to develop
Just as early evolutionary biologists looked plant varieties better adapted to improved
to breeders for empirical demonstration growing environments, with adaptation
of results of selection that illuminated measured primarily as increased yield
FIGURE 14.5
Phenotypic selection classified according to outcome of selection
Phenotypic selection for a given trait
S≈0, Phenotypic selection S>0, Phenotypic selection

is random in terms of 2P; is non-random in terms of
ecologically, genetically 2 P
and evolutionarily non-
significant
R≈0, Phenotypic selection R>0, Phenotypic selection

is random in terms of 2G, is non-random in terms of
when a) h2≈0 due to 2E 2G, because h2>0 due to
>> 2G, or, b) h2=0 high 2A cf. with 2E
because 2A =0 due to
fixation, results can be
ecologically and
agronomically significant
E≈0, Maintenance or E>0, Directional or

stabilizing selection, disruptive selection,
maintains traits between results in new populations
generations, results are & varieties, results are
genetically significant evolutionarily significant
See text for key to symbols.

(Allard, 1999: 49). Their emphasis in the ensuing proliferation of crop varieties.
selection is on achieving directional, multi- It also means formal plant breeders tend
generational, micro-evolutionary change. to judge the efficacy of farmer seed saving
This makes sense given the organization of in terms of applied evolution, i.e. the same
industrial agricultural systems (see Section criteria they apply to their own work, and
14.2.1 above, and Figure 14.3). It also means assume that farmers use these criteria as
that plant breeders often view farmers’ well.
selection of seeds or other propagules for In the following sections we describe
planting as a form of mass selection for phenotypic selection by farmers organized
heritable traits, the process that is assumed in terms of possible outcomes: longer-
to account for crop domestication and for term (multi-generational) genetic change
or micro-evolution (E) [hereafter referred of later generations, resulting from

to as ‘evolution’, in the sense of multi- limited spontaneous cross-pollination.
generational change in the context of Experimental evidence from Syria shows
agricultural crops, not in the larger biological that farmers could efficiently select among
sense of speciation]; inter-generational over 200 barley entries (fixed lines and
genetic change or response (R); and within- segregating populations), with results in
generation phenotypic differentiation (S, terms of yield potential that equalled, and
selection differential) (see Section 14.1.2 in one case exceeded, selections by plant
above, and Figure 14.1). Where possible, breeders in the same environments (Ceccarelli
we also discuss farmer goals for selection, et al., 2000). These findings indicate that
although many studies of farmer selection farmers have developed selection criteria
that document genetic or agronomic effects for identifying high yielding phenotypes
do not document farmers’ goals (and vice that are just as effective as those used by
versa). Note that, regardless of goals, the breeders, and more effective in the growing
outcomes of farmer selection can be varied, environments typical of those farmers’ own
as depicted in Figure 14.5. fields (Ceccarelli and Grando, 2007).
It is much more difficult to effect
14.5.3 Selection for evolution evolutionary change in predominantly
The clearest evidence for contemporary cross-pollinated, seed-propagated species,
farmer selection for evolution is in spe- especially for quantitative traits with low
cies that are normally propagated clon- heritability. However, as described earlier,
ally. Some Andean potato farmers search farmers can discriminate between low and
their fields for volunteer seedlings resulting high heritability traits, and use this as a
from spontaneous hybridization as a way basis for decisions about selection (Soleri et
to diversify their production (Zimmerer, al., 2002). Farmers in Oaxaca, Mexico, often
1996: 201). For example, indigenous South select maize seed with the goal of changing
American farmers intentionally incorporate or creating populations with preferred,
cassava seedlings into recognized varieties, highly heritable traits, like kernel, tassel
resulting in increased heterogeneity within and husk colours for culinary and aesthetic
varieties (Elias et al., 2001; Pujol, David reasons (e.g. maize varieties selected for
and McKey, 2005). Farmers also select the the beauty of their purple tassles) (Soleri
largest volunteer seedlings, which results in and Cleveland, 2001), while the majority
increased heterozygosity as a result of the of these same farmers see no possibility of
most heterozygous plants also being the changing the key trait of yield, which has
largest, and therefore the least likely to be low heritability, as discussed below (Soleri
eliminated during early weeding, although and Cleveland, 2001). There is evidence
farmers’ goals for this selection are unclear that farmers in central Mexico have selected
(Pujol, David and McKey, 2005). for and maintained a new landrace, based
In seed-propagated species that on seed and ear morphology, among
are predominantly self-pollinated, segregating populations resulting from the
compared with cross-pollinated species, hybridization of two existing landraces
it is relatively easy to make and maintain (Perales, Brush and Qualset, 2003). In
evolutionary changes by selecting from Rajasthan, India, there is evidence based on
among the segregating F1 plants or those research with pearl millet that farmers use
mass selection for low heritability traits in result in new varieties (i.e. the goal is E)
cross-pollinating species with the goal of (Evans, 1993: 313–314). Like plant breed-
making directional change in their varieties ers (Cooper, Spillane and Hodgkin, 2001),
(Christinck, 2002: 126; Vom Brocke et farmers also encourage gene flow under
al., 2002). This research also documented some conditions, for example mixing seed
farmers’ intentional introgression of from different sources, planting different
modern with traditional varieties of pearl populations contiguously or in same plot,
millet, and subsequent selection, resulting and by making crosses, as a way of increas-
in increased genetic variation and long- ing the variation on which to select.
term directional change (E) in selected Farmers can be successful in maintaining
traits, such as growing period (Christinck, varietal ideotypes through direct, inten-
2002: 123; vom Brocke et al., 2003a). tional selection for key traits, especially for
However, although it is clear that farmers highly heritable phenotypic traits, like those
can understand the principle of phenotypic that define a variety. This type of selection
selection and use it to achieve goals of is probably most important for cross-polli-
evolutionary change with different crops, nated crops, such as pearl millet and maize,
this may not always, or even usually, be as discussed below, since it is much more
their goal, or the result. difficult to maintain populations in these
compared with clonally propagated and
14.5.4 Selection for genetic response, self-pollinated crops. In eastern Rajasthan,
but not evolution India, amplified fragment length polymor-
Farmers also select with the goal of phism (AFLP) analysis showed that farmers
eliminating changes in phenotypic traits maintained the ideotypes of distinct intro-
resulting from gene flow or natural or duced pearl millet FVs, even though they
indirect phenotypic selection, i.e. to achieve have the same name as local FVs, via inten-
R but not E. Best documented are farmers’ tional selection of panicles for their unique
attempts to maintain varietal ideotypes based phenotypes (vom Brocke et al., 2003b). In
on quantitative or qualitative phenotypic contrast, farmers in Jalisco, Mexico, regu-
traits over time in the face of gene flow larly mix maize varieties together by clas-
(Berthaud et al., 2001). Plant breeders can sifying seed obtained from diverse sources
control unwanted gene flow much more as the same variety based on ear or kernel
effectively in their experimental plots than morphology and colour, which, together
farmers can in their fields, and in industrial with planting patterns, leads to a 1–2 per-
agriculture farmers often buy new seed cent level of gene flow between maize
every year, especially for cross-pollinated varieties during one crop cycle (Louette,
crops like maize, eliminating most concerns Charrier and Berthaud, 1997). A control-
regarding gene flow. led experiment found that, compared with
This type of farmer selection to elim- random selection, farmer selection dimin-
inate changes may contrast with main- ished the impact of gene flow on one FV
tenance (stabilizing) selection by plant from contrasting FVs for key varietal traits
breeders, which usually has the goal of (kernel rows per ear, kernel width and
maintaining yield in the face of changing kernel colour), but did not have any effect
environments by incorporating new alleles on allelic frequencies at 9 polymorphic loci
or changing allele frequencies, and may coding for traits invisible or unimportant to
farmers (Louette and Smale, 2000). Farmers or other propagules for cross-pollinated
stated that they were not interested in (e.g. in maize; Soleri and Smith, 2002), self-
changing their varieties, but in maintain- pollinated (e.g. in barley; Ceccarelli et al.,
ing varietal ideotypes, and appeared to be 2000) and vegetatively propagated crops
achieving their goal. Research in Oaxaca, (e.g. in potato; Zimmerer, 1996). Selection
Mexico, using microsatellite data support- with this goal is also conducted as part of
ed this finding in terms of the results of MV seed multiplication (Simmonds and
farmer selection, although farmers’ goals Smartt, 1999: 215). Plant breeders may also
were not investigated. Extensive gene flow carry out this type of selection, for example
and little molecular genetic structure was by removing small seed, but they do this to
observed, but the maintenance of signifi- decrease the contribution of E2 to P2 , and
cantly different maize populations based so increase heritability with the goal of E.
on morphophenological traits of interest to Research on non-heritable phenotypic
farmers persisted (Pressoir and Berthaud, differences shows these can have important
2004b). intra-generational effects in terms of ecolo-
A study in Chiapas, Mexico, found gy and agronomy. Even in species with high
that cultural diversity, as measured by heritability for seed polymorphisms, envi-
ethnolinguistic groups, was not reflected ronment may be an important determinant
in maize diversity as measured by isozyme of seed size and shape, and seed polymor-
variation, but was reflected in some phism can be a significant determinant of
morphological traits (Perales, Benz and differential survival via influence on survi-
Brush, 2005). The differences observed may vorship and adult plant size (Baskin and
have been due to unidentified culturally- Baskin, 2001: 208–214). In maize, for exam-
based networks or practices that structured ple, larger seed size was found to provide
these maize populations based on farmer significant advantages in the early stages of
selection for a few critical traits against a plant growth (from germination until stem
background of ongoing gene flow (Perales, elongation) (Bockstaller and Girardin,
Benz and Brush, 2005), as was found in 1994), and was correlated with better early
the study in the central valleys of Oaxaca, vigour, greater leaf area throughout the life
Mexico, (Pressoir and Berthaud, 2004b), cycle and more rapid development from
although neither study investigated farmer time of emergence to flowering (Pommel,
goals in detail. 1990; Revilla et al., 1999).
When the goal of selection is intra-
14.5.5 Selection for intra-generation generational phenotypic differentiation,
phenotypic difference the result may not be genetic response or
Although farmers are capable of pheno- evolution, especially for low-heritability
typic selection that is effective in achieving traits in cross-pollinated crops. This
goals of evolution and genetic response, hypothesis was supported by results of
perhaps the most common goal of farmer maize seed selection exercises with farmers
selection is not genetic, but solely pheno- in two communities in Oaxaca, Mexico.
typic, because most of the time a farmer’s The exercises were done with maize ears
primary goal in selecting seed is to obtain post-harvest, which is the way these farmers
good planting material. This often means and most others in Mexico select maize
selection for large, clean, disease-free seeds seed. Their selections resulted in high S
TABLE 14.2
Farmers’ expectations for response to selection for their primary selection criterion in the major crop
they grow
Country, crop, trait (n) Question A. Farmers responding that Question B. For farmers responding
response to intentional selection for IS10>RS10 to Question A, those stating that
10 cycles > random selection for 10 response to intentional selection for 11
cycles (IS10>RS10) cycles > random selection for 10 cycles +
intentional for 1 cycle (IS11>RS10+IS1)
n % P n % P
Mexico, maize, ear length (59) 23 39 * 0.000000 6 26 * 0.000000
Cuba, maize, ear length (29) 27 93 0.245614 12 44 * 0.000002
Syria, barley, plant height (21) 20 95 0.499999 11 55 * 0.000614
Nepal, rice, grain yield (40) 39 98 0.499999 17 44 * 0.000000
Mali, sorghum, grain yield (40) 35 88 0.057662 23 66 * 0.000078
Total (189) 144 76 * 0.000000 69 48 * 0.000000
One sided Fishers’ exact test , of the null hypothesis that, similar to plant breeders, farmers would see intentional selection
as achieving a greater response compared to random selection. Calculated using SISA (http://home.clara.net/sisa/). RS =
random phenotypic selection by farmer, IS = intentional phenotypic selection by farmer.
Based on Soleri et al., n.d.
Some farmers said large seed resulted in a hypothetical scenario asking them to
higher germination, larger seedlings, early compare random with intentional selection
vigour and higher yields, although most for 10 cycles in a typical field, in populations
farmers attributed their preference for large with phenotypic variation for the trait
seed to ‘custom’. they used as major selection criterion
It is still possible that simple mass (Figure 14.6) (Table 14.2, question A). The
selection for intra-generational phenotypic null hypothesis was that farmers did not
differences could result in R or E even if differ from plant breeders, i.e. that they
these are not farmer goals. As mentioned would all consider intentional selection to
above, it is not clear what importance this be more effective than random selection for
had during domestication and subsequent improving or at least maintaining this trait.
diversification of crops, versus intentional The majority of responses corresponded to
selection for short-term change or long- the null hypothesis of no difference between
term maintenance. For example, maize farmer and plant breeder expectations that
farmers in Uganda and the United Republic intentional selection was more effective
of Tanzania, like those in Mexico, were for increasing yield, 76.2 percent (144/189),
reported to select for large, clean kernels although those who disagreed with that
from large ears, apparently because they idea were sufficient to reject the hypothesis
believed that these germinated well and statistically (P = 0.00000). Disagreement
produced high-yielding plants (Gibson et was particularly frequent among maize
al., 2005). Interestingly, this appeared to farmers, probably due to recombination in
result in decreased resistance to maize streak that cross-pollinating crop.
virus, since resistant plants had smaller ears, These results indicate that farmers who
and plants with large ears appeared to be believe there is an advantage of intentional
non-resistant escapes. over random selection, see their goal for
As part of a comparative five-country phenotypic selection as either S or R or E.
study of FK and PBK (Soleri et al., To discriminate between these possibili-
2002, 2004), farmers were presented with ties, and with the same null hypothesis as
outlined above, those farmers responding and the movement to make formal plant
to the first question that intentional selec- breeding more relevant to farmers through
tion resulted in greater yield, were asked to PPB. Understanding farmers’ choice and
compare random selection for 10 cycles fol- selection practices, their biological results,
lowed by one cycle of intentional selection, the knowledge and goals underlying them,
with 11 consecutive cycles of intentional and the similarities and differences with
selection. Results differed significantly from plant breeders provides a means for the two
the null hypothesis (Table 14.2, question groups to work together more effectively.
B). Among these farmers, only 23.2 percent This understanding and collaboration is
(20/86) saw 11 years of intentional selection critical for supporting all of the important
as superior. These results demonstrate that functions of SSTW agriculture, including
among those farmers favouring intentional long-term global food security.
selection, only a minority see it as provid- For PPB, this means that farmers’ goals
ing cumulative multi-generational change for varietal choice and phenotypic selec-
(E), while the primary selection goal of tion need to be understood in the context
the other farmers who saw an advantage of a system that integrates production,
to multi-generational intentional selection consumption, improvement, multiplication
for low-heritability yield-related traits is and conservation. The biological result of
either eliminating changes between genera- phenotypic selection needs to be evalu-
tions (R) or a non-genetic advantage they ated in terms of its possible ecological
believe is fully achieved within one year effects (via S), as well as in terms of R
(S). The large number of farmers who do and E. Additionally, farmers’ theoretical
not consciously see an advantage to multi- knowledge of choice and selection, not just
generational intentional selection, but who, their criteria, need to be understood by
like other farmers, select for large seed from plant breeders to fully realize the potential
large, clean ears, may do so because of cus- benefits of collaboration. The value of this
tom, as did the majority of farmers in the research will be judged by its effectiveness
selection experiment described earlier. in improving the efficiency and outcomes
of collaborative breeding by scientists and
14.6 CONCLUSIONS farmers, and improvement in the well-being
Many elements of crop variety choice of those farmers and their communities.
and plant selection in the Third World
contrast substantially with industrial ACKNOWLEDGMENTS
agricultural systems, including the growing We thank the many farmers and scientists
environments, genetic resources and we have worked with in Cuba, Egypt,
organization of the agricultural system. Ghana, Guatemala, Mali, Mexico, Nepal,
The urgency of understanding farmer Pakistan, the Syrian Arab Republic and
selection will increase in the future with the United States of America for sharing
global climate changes, the continuing loss their knowledge, both about crop gen-
of genetic resources, the rapid spread of otypes and growing environments, and
transgenic crop varieties, the development their ideas about improving plant breeding
of a global system of IPR in crop genetic and crop production. Thanks to Salvatore
resources, the need to make agriculture Ceccarelli and Elcio Guimarães for com-
more sustainable while feeding more people, ments on drafts of this chapter. We grateful-
ly acknowledge the UCSB Faculty Senate, extent and implications. In [Proceedings

the US National Science Foundation (SES- of the] 7th International Symposium on
9977996, DEB-0409984), and the Wallace the Biosafety of Genetically Modified
Genetic Foundation for recent support of Organisms, Beijing, China, p. 65.
research. Al-Yassin, A., Grando, S., Kafawin, O., Tell, A.
& Ceccarelli, S. 2005. Heritability estimates
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367
CHAPTER 15
Breeding for quantitative variables

Part 2: Breeding for durable
resistance to crop pests
and diseases
Raoul A. Robinson
15.1 INTRODUCTION The switch from amateur to professional

There are three categories of plant breeder. breeding occurred because there are two
First is the professional, who is a highly kinds of breeding, depending on whether
trained scientist with a profound knowl- the breeding is for multiple-gene or single-
edge of modern genetics and related sub- gene characters. Rimpau was working with
jects. Second is the amateur, who uses multiple-gene characters, and he obtained
simple selection techniques to produce new small, quantitative improvements with each
varieties. Third is the subconscious selec- generation of selection. However, when it
tor who unwittingly changes plants by came to resistance to crop pests and diseases,
artificial selection. It is these subconscious the professional breeders were working with
selectors who were responsible for all the single-gene resistances, because they had a
original domestication of plants, and for choice and they chose this method as it gives
much of the modern artificial selection in a quick response. This single-gene breeding
Third World countries. is usually too complex and too difficult for
There were no professional plant breed- amateur breeders, and this was why they
ers before 1900, and all plant breeding was disappeared during the twentieth century.
done by amateurs and subconscious selec- Today, the pendulum is swinging back
tors. The classic example of an amateur again, and there is a new appreciation of
was a farmer called Rimpau who, in 1866, the value of both amateur breeders and
started selecting the best rye plants in his multiple-gene resistances. Participatory
crop for use as seed. He did this with each plant breeding, using multiple-gene
successive crop and, after twenty years, resistances, now provides an admirable
he had a greatly superior rye known as opportunity for cooperation between
‘Schlanstedt’, which quickly became popu- amateurs and professionals.
lar in much of Europe. In this chapter, par-
ticipatory plant breeding is taken to mean 15.1.1 Crop parasites
cooperation between professional and ama- In this chapter, a crop parasite is defined as
teur plant breeders. any organism that spends a major part of
During the twentieth century, there its life cycle inhabiting one host individual,
were major changes. Between 1900 and and obtaining nutrients from that host. The
1905, three seminal discoveries were made. host, of course, is a crop plant. A parasite
These were the recognition of Mendel’s may be an insect, mite, nematode, parasitic
laws of inheritance, Johannsen’s discovery angiosperm (e.g. broomrape, witchweed,
of pure lines, and Biffin’s discovery of dodder), or any of the various categories of
single-gene resistances. All subsequent plant pathogen, such as fungi, bacteria and
plant breeding was done by scientific viruses. However, weeds are not parasites:
professionals, who were totally captivated they are competitors, and they are not part
by these new discoveries. Amateur plant of this discussion.
breeders disappeared almost entirely. This chapter is addressed to both partic-
More recently, the techniques of genetic ipating, amateur plant breeders, who may
engineering have become popular, but find Return to Resistance (Robinson, 1996)
these, of necessity, involve single-gene helpful, and to cooperating professional
resistances, and they usually require breeders who may find Self-Organising
professional scientists. Agro-Ecosystems (Robinson, 2004) useful.
Breeding for quantitative variables. Part 2: Breeding for durable resistance to crop pests 369
15.1.2 Plant breeding break down as new races, strains, biotypes

For the whole of the twentieth century, or pathotypes of the parasite emerge.
scientific plant breeding has had four main Horizontal resistances provide stable
objectives. These were improvements protection. That is, they are beyond the
in (i) yield; (ii) quality of crop product; capacity for micro-evolutionary change of
(iii) agronomic suitability; and (iv) resistance the parasite, which is consequently unable
to parasites. This scientific breeding has to produce a new strain that is unaffected
been remarkably successful in the first three by that resistance. Other protection mecha-
of these objectives, but much less successful nisms can also be stable. Examples of sta-
with breeding for resistance. The basic ble insecticides include natural pyrethrins,
reason for this has been that resistance kept rotenone, and a film of oil on water to
breaking down because of new strains of control mosquito larvae. Examples of stable
the parasite. There was then an apparently fungicides include both copper and dithio-
endless repetition of resistance failures carbamate formulations.
and a ‘boom and bust’ cycle of cultivar
production. 15.1.4 Vertical resistance
Vanderplank (1963, 1968) first made a Vertical resistance has several remarkable
clear distinction between single-gene and advantages. First, it provides complete
multiple-gene resistances to crop parasites. protection against a parasite. There are a few
He called single-gene resistance ‘vertical examples of incomplete vertical resistance
resistance’, and it is normally qualitative (see below) but not enough to invalidate
in that it provides complete protection this rule. Second, it usually has a very wide
or none at all, with no intermediates. He climatic range and can be employed across
called multiple-gene (polygenic) resistance broad geographical regions. Third, being
‘horizontal resistance’, and it is quantitative controlled by single genes, it is amenable to
in that it can provide every degree of pro- gene-transfer and back-crossing techniques.
tection from a minimum to a maximum. These techniques are so elegant, and so
beautiful, that they were greatly favoured by
15.1.3 Stability and instability professional breeders during the twentieth
Any protection mechanism against a crop century.
parasite may be described as unstable or However, vertical resistance does have
stable. An unstable resistance is within the some grave disadvantages. First, as already
capacity for micro-evolutionary change of mentioned, it is unstable. It is liable to
the parasite. This means that the parasite is break down when faced by new strains
able to produce a new strain that is unaf- of the parasite. The speed of this break-
fected by that protection, which is then said down can vary greatly. The fastest occurs
to have ‘broken down’ (strictly speaking, in the first growing season, and this hap-
the protection is unaltered, and it is the pened with Puccinia polysora of maize in
parasite that has changed). Many synthetic tropical Africa (see below), and in Late
insecticides and fungicides provide unstable potato blight (Phytophthora infestans) in
protection, and they sooner or later break the Toluca Valley of Mexico, which is the
down in the face of new strains of the centre of origin of this fungus. The slowest
parasite. Single-gene, vertical resistances breakdowns take so long to occur that the
are almost always unstable, and they too vertical resistance is effectively durable (see
below), but these are too rare to be a gen- The main advantage of horizontal resist-
eral breeding tool. ance is that it is durable, and that it is
A second disadvantage of vertical possible to breed for increased levels of
resistance is that it is responsible for the many different quantitative variables simul-
vertifolia effect, which is the gradual loss of taneously. Participatory plant breeders can
horizontal resistance during breeding for accordingly aim at high levels of horizontal
vertical resistance, and which is described resistance to all locally important para-
in more detail below. A third disadvantage sites. This will achieve crop husbandry that
is that vertical resistances occur only against is effectively free of all parasite damage,
some species of parasite. Consequently, it is and one that is free of pesticides as well.
not possible to use vertical resistance for all And these freedoms will be permanent.
the locally important parasites. However, it must be emphasized that this is
In general, vertical resistance is not the objective. Such an objective may prove
recommended for participatory plant impossible to achieve in practice, at least in
breeding. This is mainly because the failure some crops, and in some areas. But, even if
of a wonderful new cultivar, which has this objective is unattainable, there will be at
taken years of devoted work to produce, least some improvement over current farm-
by both professional and amateur breeders, ing practices in terms of increased yields
is quite frankly heart-breaking. Nothing and decreased damage from parasites.
can be expected to discourage amateur
breeders more than this. An essential aspect 15.1.6 Two kinds of plant breeding
of participatory plant breeding is that we Clearly, the key difference in breeding crop
maintain the confidence of the participating plants for temporary and durable resist-
amateurs. Consequently, participating ances is the difference between breeding for
professional breeders should be very single-gene and multiple-gene characters.
cautious about recommending the use of Breeding for single-gene characters
vertical resistance, or even the combined requires both pedigree breeding and back-
use of vertical and horizontal resistance, in crossing, or the very modern techniques of
participatory plant breeding. marker assisted selection and genetic engi-
neering. Anyone using these techniques for
15.1.5 Horizontal resistance acquiring resistance should assume that the
Being a quantitative variable, horizontal resulting resistance will be unstable, and
resistance can be expressed at any level that it will have a very high probability of
between a minimum and a maximum. In the breaking down sooner or later.
absence of crop protection chemicals, the Breeding for multiple-gene characters,
minimum level of horizontal resistance usu- such as horizontal resistance, requires an
ally results in a total loss of crop from the entirely different breeding technique, called
parasites. And the maximum level of hori- recurrent mass selection, which is discussed
zontal resistance usually results in negligible below.
loss of crop. However, the maximum level of
horizontal resistance never provides as com- 15.2 WHY WAS TEMPORARY
plete a protection as vertical resistance. Even RESISTANCE SO POPULAR?
with the highest level of horizontal resist- During most of the twentieth century, ver-
ance, there is always some slight parasitism. tical resistance was consistently the resist-
ance of choice, and horizontal resistance Being quantitative, in contrast, horizontal

was almost entirely ignored. This is a his- resistance is easily obscured, its durability
torical fact, and it is so important that we not recognized, and its value consistently
must examine its causes in some detail. underestimated. A clear comprehension of
The effectiveness of horizontal resistance these misleading variables, these sources of
is influenced by many quantitative variables. error, is consequently essential for anyone
Many of these variables are difficult to wishing to breed crops for durable resist-
observe or measure, and there is a powerful ance. The twelve most important of them
tendency to neglect them. Twentieth- are considered below. They are summa-
century plant breeders did neglect them, and rized in Table 15.1.
this is why they also neglected horizontal
resistance. They concentrated on vertical 15.3.1 Parasite interference
resistance because it is complete. This The only way to measure the level of
completeness was very attractive. It meant horizontal resistance is by the level of
that the parasite control was total. However, parasitism. Parasite interference can make
the ephemeral nature of the resistance was such assessments wildly inaccurate, and
usually revealed only much later. the level of parasitism is then a thoroughly
Today, if we are to breed crops misleading indication of the level of
successfully for resistance that is durable, horizontal resistance. Parasite interference
we must understand these misleading, occurs because the parasites are mobile,
quantitative variables that disguise the and they can move from plot to plot, or
effectiveness of horizontal resistance. from plant to plant. The importance of
Within the framework of participatory this phenomenon was first discovered by
plant breeding, a primary function of the Vanderplank (1963).
participating specialists must be to ensure Parasite interference can be seen with field
that the participating farmers are not trials in which the parasites can move from
deceived by these misleading variables. one plot to another. The results of parasite
interference can be extraordinary, and
15.3 MISLEADING VARIABLES may lead to errors of several hundred-fold
In principle, breeding for horizontal resist- (James et al., 1973). In these circumstances,
ance is very simple and very easy and for the effects of horizontal resistance can be
this reason it is ideal for participatory plant totally obscured. Parasite interference is
breeding. But there are a number of factors at its worst in the very small screening
that can be horribly misleading and which, plots produced by the technique of ‘ear-to-
unless understood, can lead to totally false row’ selection (or family selection), often
observations. These false observations often used during the breeding of self-pollinating
disguise horizontal resistance so effectively small-grain cereals and pulses. The parasite
that little genetic advance can be seen. interference can be so intense that plants
It was these sources of error that led with a functioning vertical resistance can
to vertical resistance being the preferred appear diseased, because of millions of
resistance mode among plant breeders for hypersensitive flecks produced by the
the whole of the twentieth century. Being failed infections of non-matching strains of
qualitative, vertical resistance is easily seen, the parasite. Imagine the level of parasitism,
but its unstable nature is not apparent. therefore, if those infections are produced
TABLE 15.1
Summary of misleading variables
Misleading Variable Problem Solution
Parasite interference Hides high levels of horizontal resistance Use relative measurements only
Epidemiological Resistance requirements vary between agro- Use on-site screening
competence ecosystems
Environmental erosion An apparent loss of horizontal resistance Each agro-ecosystem should have its own
due to re-location in an area of higher breeding programmes
epidemiological competence
Vertifolia effect The loss of horizontal resistance when Inactivate all vertical resistances, and avoid
breeding crops for vertical resistance, or any use of crop protection chemicals
under the protection of crop protection
chemicals
Parasite erosion An apparent loss of horizontal resistance Screen in an area of high parasite density
due to changes in the parasite population
False erosion An apparent loss of horizontal resistance Be more careful
due to sloppy or negligent assessments
Biological anarchy An increased epidemiological competence in This phenomenon will disappear after a few
the parasite due to the loss or debilitation seasons of freedom from pesticides, as the
of biological control agents biological controls are restored by the use of
horizontal resistance
Population immunity Makes field assessments preferable Avoid laboratory assessments of resistance
Chance escape Provides a false indication of resistance Inoculate the screening population, or use
grid screening
Quantitative vertical Looks like horizontal resistance but is not Use parents that lack vertical resistance
resistance genes, or use the one-pathotype technique
(see below).
Durable vertical This rare phenomenon provides a false hope Do not rely on this remote possibility
resistance for breeders of single-gene resistances
Sub-optimization Do not breed for a single resistance Use the holistic approach (see below)
mechanism, or resistance to single species of
parasite
by a matching strain, against which the The misleading effects of parasite

vertical resistance does not operate. interference can be avoided during breeding
Parasite interference can also occur for horizontal resistance by using relative
among individual host plants. Consider a measurements during screening. That is,
screening population in which there are wide the least parasitized individuals are selected
differences in horizontal resistance between regardless of how severely parasitized they
individual plants. A quantitatively resistant might be. In the early breeding cycles, these
plant may be surrounded by susceptible least-parasitized individuals may well look
plants. The overcrowded parasites will then so awful that the breeder could be forgiven
move onto that resistant plant, making it for concluding that there is no point in
look far more susceptible than it really is. continuing. But that would be a mistake.
Once again, the magnitude of these errors However awful they may look, they are
can be great. In fact, if that resistant plant the least susceptible individuals of an early
were growing in a farmer’s field, as a pure breeding cycle and, as such, they should
line or clone, there would be no parasite become the parents of the next breeding
interference, and its resistance might then cycle.
be entirely adequate to provide complete Remember also that we are breeding
control of the parasite. for comprehensive horizontal resistance
to all the locally important parasites. This epidemiological competence is low, the
requires the one quality of ‘good health’. horizontal resistance will also be low,
The least parasitized individuals will be because there is little selection pressure for
fairly susceptible to many different species resistance. But where the epidemiological
of parasite. Their quality of ‘good health’ competence is high, the level of horizontal
will be low. But it will be higher than all resistance will also be high, because there is
those other, less healthy individuals in the strong selection pressure for resistance.
screening population, many of which may When breeding for horizontal
have disappeared entirely. resistance, this variation in epidemiological
A final comment about parasite interfer- competence is important in two ways.
ence concerns the movement of parasites First, we must use ‘on-site screening’. This
from one farm, or one district, to another. means that the screening for horizontal
Many organic farmers are able to culti- resistance must be conducted: (i) in the
vate fairly susceptible cultivars successfully locality of future cultivation; (ii) in the time
because their neighbours are using crop of year of future cultivation; (iii) in the field
protection chemicals. The district interfer- (i.e. not in the laboratory or greenhouse);
ence is then minimal. But were all the farm- and (iv) according the farming system (e.g.
ers in that district to eschew crop protection organic or conventional, irrigated or rainfed)
chemicals, the parasite populations would of future cultivation. On-site screening is
be so large, and the district interference particularly well suited to participatory
so great, that the use of those cultivars for plant breeding.
organic agriculture might prove impossible. Second, each distinct agro-ecosystem
The other side of this coin is that progress will require its own horizontal resistance
in breeding for horizontal resistance will breeding programme for each of its crop
lead to reductions in district interference, species. This programme must be aimed at
which, in turn, will enhance the effects of the levels of epidemiological competence of
that horizontal resistance. the locally important parasites. In practice,
this is not difficult. Agro-ecosystems are
15.3.2 Epidemiological Competence usually quite large, and the necessary levels
Epidemiological competence refers to the of horizontal resistance will be discovered
ability of a crop parasite to cause an epi- by practical farming experience, spread
demic (or infestation). It is another bio- over time, as the breeding programmes
logical variable that can be expressed at any produce more and more new cultivars, with
level between a minimum and a maximum. higher and higher levels of comprehensive
Consider a wild ecosystem, which horizontal resistance.
might, perhaps, extend up a mountainside.
The epidemiological competence of a plant 15.3.3 Environmental erosion of
parasite might change along a gradient horizontal resistance
from low to high moisture, or temperature, If a cultivar has adequate horizontal resist-
or whatever factor is governing that ance in an agro-ecosystem in which the
epidemiological competence within the parasite has a relatively low epidemiological
ecosystem. The various host ecotypes competence, and that cultivar is taken to a
along that gradient will have corresponding different agro-ecosystem, where the parasite
levels of horizontal resistance. Where the has a high epidemiological competence, the
resistance will appear to have decreased, cannot be observed or assessed. Because

possibly disastrously. This is known as envi- individuals with the highest levels of
ronmental erosion of horizontal resistance. horizontal resistance are always a minority
Strictly speaking, of course, the resistance is in a screening population, less-resistant
unchanged and it is the epidemiological com- individuals are more likely to be selected on
petence of the parasite that is different. But the basis of their other attributes.
the level of parasitism increases and this can After many generations of crop
be alarming if the cause is not understood. breeding conducted under the protection of
It is because of these differences in pesticides or vertical resistance, or both, the
epidemiological competence between levels of horizontal resistance in modern
agro-ecosystems that we speak of ‘locally cultivars are usually low. This erosion of
important parasites’. We are breeding agro- horizontal resistance has continued for
ecotypes (i.e. cultivars) for our own agro- about a century in many species of crop.
ecosystem. These agro-ecotypes may have It has been particularly serious in potatoes,
too much, or too little, resistance in other tomatoes and cotton, but there are few
agro-ecosystems, where the epidemiological species in which it has not occurred. It is
competences are different. But these other also the reason why breeding for horizontal
agro-ecosystems are not our concern. resistance is now so important.
Equally, other people’s agro-ecotypes will The lesson of the vertifolia effect is
very likely prove inferior in our own agro- that we must never protect our screening
ecosystem. Professional breeders often speak populations with crop protection chemicals.
of site-specific plant breeding in which the However, there is one exception to this
breeding targets are aimed at a particular site rule. In the early screening generations,
with special requirements. On-site selection even the least parasitized individuals may
fits in well with this concept. be so severely damaged that their very
The environmental erosion of horizon- survival is threatened. In such a case it is
tal resistance seems like a breakdown of permissible to use crop protection chemicals
that resistance but, clearly, it is not. as a last resort to preserve the parents of the
next generation. Equally, we must never
15.3.4 The vertifolia effect use vertical resistances in our screening
The vertifolia effect was first recognised populations. Methods of avoiding this are
by Vanderplank (1963) and it is an ero- described below.
sion of horizontal resistance resulting from People who call themselves ‘seed sav-
genetic changes in the host species which ers’ know that century-old cultivars, often
occur during breeding for vertical resist- called ‘heirloom varieties’, usually have
ance. These changes can also occur during higher levels of durable resistance than
any breeding that is conducted under the modern cultivars. This difference is a meas-
protection of crop protection chemicals. ure of the overall vertifolia effect that has
This erosion occurs because horizontal occurred during the twentieth century.
resistance can be measured only in
terms of the level of parasitism. If the 15.3.5 Parasite-erosion of horizontal
level of parasitism is totally obscured by resistance
pesticides, or by a functioning vertical Occasionally there can be changes in
resistance, the level of horizontal resistance the parasite population that lead to an
increased epidemiological competence. 15.3.7 Biological anarchy

This can happen, for example, with the Biological anarchy is the converse
Fusarium and Verticillium wilt fungi. The of biological control. It means that the
frequency of the highly pathogenic forms various agents of biological control have
of these soil-borne pathogens might be been depleted or destroyed by crop
quite low in the area of field screening. protection chemicals, particularly when
However, with repeated use of one field there has been pesticide overload. These
for the screening work, the frequency of agents might be hyper-parasites, predators,
pathogenic forms will increase, more or microbiological competitors, toxin-
less in step with the increases in horizontal producing micro-organisms, or organisms
resistance in the host. The overall effect is that stimulate resistance responses in the
then an appearance of no progress whatever, host. The importance of biological anarchy
with a very real possibility of the breeding is revealed by the success of integrated pest
programme being abandoned. management, generally known as IPM. This
This is a somewhat rare phenomenon method involves careful monitoring of the
and the participatory breeder is unlikely to crop parasites in order to reduce the use of
be faced with it very often. However, it is crop protection chemicals to the absolute
as well to be aware of the possibility, just in minimum necessary for control. A gradual
case of apparently discouraging progress. increase in biological control then occurs.
As a general rule, however, the parasitic When the biological controls are restored,
ability of most crop parasites, particularly a greatly reduced rate of crop protection
the obligate parasites, is fixed, and parasite- chemical application can be maintained.
erosion is rare. The importance of biological anarchy
during breeding for horizontal resistance is
15.3.6 False erosion of horizontal that, in the absence of biological controls,
resistance many crop parasites will behave with
There can be false erosion of horizontal a savagery that would be impossible if
resistance due to sloppy techniques, or just the biological controls were functioning.
plain carelessness. The classic example of This means that assessment of the level
this occurred with a virus disease of sugar of horizontal resistance can be difficult.
cane called mosaic. This disease can be Screening for horizontal resistance
devastating when susceptible cultivars are should be conducted in an area where
being cultivated. However, in most areas, there is no biological anarchy. But, given
the disease was controlled so completely the widespread use of crop protection
with horizontal resistance that it tended chemicals, particularly in the industrial
to be forgotten. New cultivars that were nations, it is often impossible to find
released to farmers were susceptible such an area. The screening population
because they had been inadequately tested, then appears to have so little resistance
or not tested at all, for resistance to this that grave doubts develop concerning the
virus, and there would then be a flare wisdom of this approach. However, the
up of the disease. It was not uncommon problem of biological anarchy is less acute
for this to be blamed on a breakdown of in developing countries, where the use of
resistance when, of course, it was nothing crop protection chemicals is much less
of the kind. intense.
It should be added that the best way Indeed, it must be strongly positive if it is
to restore biological controls is to use to become a serious crop pest or disease.
horizontal resistance. And the best way to Each individual parasite must spawn more
enhance the effects of horizontal resistance than one new individual before it dies. Now
is to restore biological controls. The two suppose that the combination of horizontal
effects are mutually reinforcing. The resistance and biological controls is such
practical effect of this is that a new cultivar that, on average, each parasite individual
with apparently inadequate horizontal spawns less than one new individual. The
resistance may well prove to have adequate parasite population growth is now negative,
horizontal resistance, once the biological even though the host individuals are not
controls are restored. This restoration immune. This situation is population
may require several seasons of cultivation immunity, and it is important in horizontal
without pesticides but, once complete, the resistance breeding in three quite different
effects can be dramatic. ways.
It should also be added that the agents First, population immunity means that
for biological control often depend on a we do not need to breed for the maximum
small population of the parasite in order levels of horizontal resistance. We need to
to maintain their own populations. A low breed for enough horizontal resistance to
level of parasitism is often desirable for this cause population immunity, and no more.
reason, provided that it has no deleterious This level is discovered from practical
effect on the yield or quality of the crop farming experience.
product. However, purchasers of organic Second, although vertical resistance can
food often like to see minor parasite be assessed on detached leaves in a test tube,
damage as evidence for freedom from crop or leaf disks in a Petri dish, or even entire
protection chemicals. plants in a growth chamber, horizontal
resistance should not be measured in this
15.3.8 Population immunity way. This is because these laboratory
Population immunity means that a host methods cannot possibly represent the
population is effectively immune, even effects of biological controls or population
though the individuals that make up that immunity. The levels of horizontal
population are less than immune. This resistance are best determined in the field
is because population growth, unlike an and, if at all possible, under conditions of
individual’s growth, can be positive or restored biological controls. Once again,
negative. Positive population growth means the best determinations will be the result of
that there are more births than deaths, practical farming experience.
and the population is increasing. Negative Third, all measurements and descriptions
population growth means that there are of horizontal resistance must be relative.
more deaths than births, and the population There can be no absolute measurements.
is decreasing. The dividing line occurs when We can describe a new cultivar “A” as being
the births and deaths are equal, and the more resistant to a particular parasite than
population growth is then zero. cultivar “B”, but less resistant than cultivar
Now consider a crop parasite. In order “C”. But we cannot have an absolute scale
to cause an epidemic (or infestation), the of measurement comparable to the Celsius
parasite population growth must be positive. scale of temperatures. This is because these
biological variables are too imprecise to of parasite and, with participatory plant
define accurately. However, this does not breeding, this will be the responsibility of
make horizontal resistance any less useful the professionals.
in farmers’ fields. If the parasite is wind-borne, such as most
fungal spores, or a flying insect, a previously
15.3.9 Chance escape prepared population of the parasite can be
The distribution of parasites in a screening released, blown or water-sprayed on to
population is often uneven. This is called a the screening population. Once again, the
‘patchy distribution’ and it is most common details of the techniques vary and will be
with soil-borne parasites and gregarious handled by the professionals.
insect pests. There may then be some host With some parasites, inoculation is not
individuals that have no parasites at all, feasible for technical reasons. An alternative
and they give the impression of being technique is then to ignore those parts of
highly resistant. They are known as ‘chance the screening population that are free of
escapes’ and, obviously, they should not the parasite in question. Or any individual
be selected as parents of the next screening plant that is entirely free of the parasite in
generation because they could be very question can be ignored. This procedure
susceptible. But the problem is how do runs the risk of discarding some highly
we recognise them, and how do we avoid resistant potential parents, but this wastage
them? There are a number of techniques is preferable to the risk of selecting highly
that increase the accuracy of the screening, susceptible escapes as parents of the next
depending on the nature of the parasite. generation.
It is in this area that the professionals will Finally, there may be parasite gradients
be most useful to participatory amateur within the screening population in which
breeders. the intensity of parasitism changes
If the parasite is soil-borne, such as a gradually from low to high from one part
root nematode, or a fungal or bacterial of the host population to another. This
wilt organism, it is a good idea to pre- effect can be eliminated by dividing the
germinate the seedlings in flats or peat screening population into a grid of suitably
pots for later transplanting in the field. sized squares. Only the least parasitized
These flats or pots can then be inoculated individual is selected within each square,
with the parasites in question, and the very regardless of the level of that parasitism
process of transplanting will ensure an even when compared with other squares.
distribution of the parasite.
If the parasite is a gregarious insect, in 15.3.10 Quantitative vertical resistance
which all individuals tend to congregate Occasionally, vertical resistance can be
on one host plant, they can often be quantitative and it is then easily confused with
redistributed on a daily basis by disturbing horizontal resistance. It occurs, for example,
them. This can be particularly important with Hessian fly (Mayetiola destructor)
with virus vectors. of wheat (Dent, 1998). Fortunately, this
If the parasite is seed-borne, it is usually situation is rare and need not worry the
feasible to inoculate the seed before sowing. breeders of most crops. However, if it does
The details of the techniques for doing occur, quantitative vertical resistance must
this vary considerably with different kinds obviously be avoided or inactivated during
breeding for horizontal resistance. This can 15.3.12 Sub-optimization

be done either by using only parents that are Sub-optimization means emphasizing some
known to possess no genes for quantitative subsystems at the expense of others, usually
vertical resistance. Alternatively, the ‘one- by working at too low a systems level.
pathotype technique’ (described below) There are many levels of subsystem in
may be employed. biological systems and sub-optimization
can lead to two kinds of error. First, by
15.3.11 Durable vertical resistance working at too low a systems level, other
Very occasionally, single-gene vertical subsystems may be overlooked. Second,
resistances may be durable. This happens, emergent properties, which can be observed
for example, with cabbage yellows only at their own systems levels, may also
(Fusarium oxysporum f.sp. conglutinans) be overlooked.
in the United States of America, potato An obvious example of sub-optimiza-
wart disease (Synchytrium endobioticum) tion in biology occurs with the ‘schooling’
in Britain, and wheat stem rust (Puccinia of fish. This schooling is an emergent prop-
graminis) in Canada (Vanderplank, 1978). erty in which a population of fish swim as
The durability is usually due to local one individual. A scientist studying only
circumstances that would not occur in one fish, or even a pair of mating fish, in
the wild pathosystem of the host and an aquarium, cannot possibly observe this
parasite progenitors. However, durable emergent property of schooling. In order to
vertical resistance can be very useful, and study schooling, the scientist must work at
there is no reason why it should not be the higher systems level of the population.
exploited. But durable vertical resistance During the twentieth century, plant
is so rare that it should not be aimed breeding for resistance to parasites suffered
at, nor depended on, in most breeding considerably from sub-optimization.
programmes. It led many plant breeders Research was conducted at the systems
astray in the twentieth century, as they level of the individual host or parasite, or
continued to hope, over-optimistically, even the individual resistance mechanism.
that their single-gene resistances would It should also have been conducted at the
also prove to be durable. level of the two interacting populations of
A technique that has had some success is host and parasite, which is a systems level
the ‘pyramiding’ of single-gene resistances. now called the pathosystem. A pathosystem
This involves putting as many different is a subsystem of an ecosystem and, when
vertical resistance genes as possible into host resistance is studied at this level, very
one plant in a so-called ‘pyramid’, and this different pictures of both vertical resistance
can prolong the life of vertical resistance, and horizontal resistance emerge. This is
particularly if genes from different species because of previously unobserved emergent
of wild plants are combined. However, this properties.
is resistance breeding at its most difficult We now recognize the significance of
and is of doubtful value for participatory two different kinds of infection. Infection is
plant breeders. If it is employed, it will defined as the contact made by one parasite
require professional plant breeding at a individual with one host individual for
plant breeding station combined with purposes of parasitism. With allo-infection,
selection in farmers’ fields. the parasite has to travel to its host, having
originated somewhere else. This is analogous the lock of the host and, consequently, all
to cross-pollination, or allogamy. With auto-infection is matching infection. Even
auto-infection, the parasite is born, hatched parasites that reproduce sexually, such as
or spawned on the host that it is infecting. many insects, will soon reach homozygosity
This is analogous to self-pollination, or of the matching biotype. However, vertical
autogamy. We also recognize that, in a wild resistances against insects are rather rare,
pathosystem, vertical resistance can control and this may explain why there has been so
allo-infection only, and auto-infection can little crop breeding for resistance to insects
be controlled only by horizontal resistance pests.
(although horizontal resistance can also These functions of the two kinds of
control allo-infection). resistance are emergent properties that
The control of allo-infection by ver- were completely unknown until recently,
tical resistance apparently operates as a and, being unknown, they were inevitably
system of locking, with each host having ignored by crop scientists during the
a biochemical lock, consisting of several twentieth century.
resistance genes, and each parasite having a Another obvious example of sub-
biochemical key consisting of several para- optimization occurs when breeding for a
sitism genes. If the parasite key does not fit single resistance mechanism, such as hairy
the lock of the host it is allo-infecting, the leaves that repel an insect pest. Horizontal
infection fails, while if the key does fit the resistance to insects usually consists of
lock the infection succeeds. Such a system many obscure mechanisms, all of which may
ensures that the frequency of matching vary quantitatively, and which collectively
allo-infection is low, and this stabilizes the reduce the rate of population growth of
population explosion of the parasite. This that pest.
stabilization is an emergent property that The converse of sub-optimization is
can be seen only at the level of the system known as the holistic approach, which
of locking, the level of the pathosystem. leads to local optimization of all variables.
However, if every door in the town has When breeding for horizontal resistance,
the same lock, and every householder has therefore, we must not sub-optimize. We
the same key, which fits every lock, the must work at the systems level of the agro-
system of locking is ruined by uniformity. ecosystem. Within this agro-ecosystem, we
And this is exactly what we have done in must use population breeding to produce
agriculture, with our use of a single vertical adequate and durable resistance to all locally
resistance in a uniform pure line, clone or important species of parasite, by exerting
hybrid cultivar that might be grown over selection pressure for the one characteristic
a huge area as a homogeneous population. of ‘good health’. Susceptibility to only one
This was sub-optimization at its worst. important species of parasite will result in an
Auto-infection can be controlled only inferior cultivar, and this would constitute a
by horizontal resistance because auto- clear case of sub-optimization. In addition
infection can commence only after there has to ‘good health’, new cultivars should have
been a matching allo-infection. Many crop good levels of all the other variable attributes
parasites reproduce asexually to produce necessary to a productive agriculture. This
a clone. All the individuals within that approach does not necessitate participatory
clone have the same key which matches plant breeding, and some professional
breeders may choose to work on their own This crop is called corn in North America
in a scientific institution. However, this but it is called maize in all other countries,
approach does depend on high numbers of and in all other languages. It was taken by
plants being screened, and many amateur the Spanish from the New World to the
breeders, working cooperatively with Iberian peninsula some five centuries ago
a professional, can lead to both greatly and tropical rust was either left behind or
increased numbers of plants screened, and it failed to survive outside the tropics. The
greatly increased attention applied to each Portuguese then took rust-free maize to
plant screened. Africa and all points east, where it was cul-
tivated for more than four centuries in the
15.4 BREEDING CROPS FOR DURABLE absence of tropical rust. This negative selec-
(HORIZONTAL) RESISTANCE tion pressure led to the level of horizontal
In this section there are inevitable resistance to tropical rust declining to its
generalizations that do not apply to all minimum natural level. In technical terms,
crop species. For example, comments about this is the Hardy-Weinberg equilibrium.
open-pollinated crops may not apply to self- In theory, it should be possible to breed
pollinated crops, or comments about annual experimentally for absolute susceptibility,
crops may not apply to perennial crops. but this is a somewhat academic point.
When planning a breeding programme, With the development of trans-Atlantic
therefore, readers should highlight only air transport in the 1940s, tropical rust was
those aspects of these descriptions that accidentally taken from the new world to
apply to their crop species of choice. the old. Devastating epidemics developed
in the low altitude, equatorial tropics. In
15.4.1 Maize in tropical Africa East Africa, a classic breeding programme
The best way to breed for horizontal for vertical resistance was initiated. Genes
resistance is to imitate the behaviour of for resistance had to be imported from
open-pollinated maize (Zea mays) in tropical America because none could
tropical Africa, following the introduction be found in the local maize populations
of the re-encounter disease called Tropical (Storey et al., 1958). Unfortunately, these
rust (Puccinia polysora). A re-encounter vertical resistances broke down so quickly
parasite is one in which the host was that none lasted long enough to be released
separated from its parasite and taken to farmers.
to another part of the world. At a later However, the appearance of this
date, the parasite is also taken to that re-encounter disease exerted positive
new area where it re-encounters its host, selection pressure for horizontal resistance
which has lost resistance in the meanwhile. in the farmers’ open-pollinated crops. After
Conversely, a new encounter parasite is about a dozen maize generations, the levels
one which evolved separately from its host, of disease had declined from ‘total loss
on a botanical relative. Later the two are of crop’ to ‘negligible loss of crop’. The
brought together in a new encounter. An horizontal resistance had increased from its
old encounter parasite is one in which the minimum level to its maximum level. With
host and parasite have never been separated, two crops each year, this transformation
even though both may have been moved to occurred in about six years, and it had
new areas (Buddenhagen, 1987). happened without any help from plant
breeders or plant pathologists. It is common leaf blight of rubber, caused by Microcyclus

knowledge that this horizontal resistance ulei. However, coffee is functionally
has now endured for half a century without deciduous in the dry season with respect
any suggestion of breaking down to new to rusted leaves only, and rubber is a
strains of the parasite. deciduous species in spite of growing in
the Amazon valley, which is permanently
15.4.2 The inactivation of vertical warm and wet.
resistance The second method involves inactivation
It is impossible to breed for horizontal of any vertical resistance genes that may be
resistance if the screening population is present in the screening population. This is
protected by functioning vertical resistanc- necessary in crops, such as wheat, in which
es. In other words, it is possible to breed for it is extremely difficult, if not impossible, to
horizontal resistance only after the vertical find parents with no vertical resistance genes.
resistances have been matched. This inactivation can be achieved with the
There are several methods of ensuring ‘one-pathotype technique’. The first step is
that no vertical resistances are functioning to designate a single vertical pathotype of
during screening for horizontal resistance. the parasite in question. Potential parents
The first method is to use only parents that are then screened for susceptibility to this
possess no genes for vertical resistance. This designated vertical pathotype. Only those
is usually possible only in crops species lines that are susceptible to the designated
that have had foreign vertical resistance vertical pathotype can become original
genes inserted into them. For example, parents of the screening population. All
the vertical resistance genes to Late potato recombinations of the vertical resistance
blight (Phytophthora infestans) were genes that occur in all subsequent breeding
inserted into cultivated potatoes from the generations will then be matched by the
wild Solanum demissum. designated vertical pathotype, which is used
In this context, it is important to note to inoculate each screening population.
that vertical resistance genes occur only Amateur breeders will have difficulties
in seasonal tissues in which there is a with this somewhat complex but essential
discontinuous pathosystem. That is, in each procedure, and in a participatory breeding
new season, each host individual is free of programme it should be the responsibility
the parasite and must be newly allo-infected. of the breeder or pathologist. The details of
This situation is seen in the seasonal tissues the technique have been described elsewhere
of annual plants, or the leaves and fruits (Robinson, 1996, 2004).
of deciduous trees and shrubs. Vertical A third possibility is to rely on natural
resistances can also occur in the seasonal, matching of any vertical resistances that
aerial tissues of perennial crops such as may be present in the screening population.
hops and potatoes. But vertical resistances Any individual that has no parasitism is
are not found in perennial crops such as inspected for the presence of the necrotic
sugar cane, cassava and sweet potato, or in spots that are typical of a vertical resistance
evergreen tree crops such as olives, citrus, reaction, and affected individuals are
tea and cocoa. Apparent exceptions to this discarded. However, this method can be
rule occur in Coffee leaf rust caused by very wasteful of breeding material if there
Hemileia vastatrix, and South American are many such plants.
It is not clear whether the maize in and with each breeding cycle the levels
tropical Africa lacked vertical resistance of horizontal resistance increase until no
genes, or that it had such genes but their further increase is either possible or neces-
resistances were matched so quickly that sary. This process of quantitative increase,
they went unobserved. In either event, the in which the progeny have a higher level of
positive selection pressure for horizontal a quantitative variable than their parents, is
resistance was not hindered by functioning known as transgressive segregation.
vertical resistance. So, when breeding for horizontal resist-
ance, there is no need to begin with a good
15.4.3 Genetic resources source of resistance, but there must be a
It is now a plant breeding shibboleth that reasonably wide genetic base to ensure that
breeding for resistance requires a ‘good all the necessary polygenes are present.
source’ of resistance before the breeding In practice, it is much easier to breed
can even begin. This is true when breeding for horizontal resistance than it is to
for vertical resistance, because it is essential breed for high yield, high quality of crop
to have at least one gene for resistance. But product or high agronomic suitability.
it is not true when breeding for horizontal It is therefore best to use high-yielding,
resistance. This point is so important that it high-quality, agronomically suitable, but
merits careful explanation. susceptible, modern cultivars as the original
Consider a heterogeneous screening parents. With suitable selection procedures,
population in which every individual is it should be easy to gradually increase
different from every other individual, but the levels of horizontal resistance, while
in which all the plants are susceptible. Each retaining the other desirable qualities.
plant possesses about 10 percent of the total Conversely, it would be very difficult to use
polygenes that contribute to the horizontal highly resistant, primitive archetypes as the
resistances to each of the various locally original parents, and then try to improve
important parasites. The host population as their various agricultural attributes, while
a whole is thus very susceptible. However, retaining their resistance.
we may assume that each host individual The maize of tropical Africa illustrat-
possesses a different 10 percent of those ed this point conclusively. The horizontal
total polygenes. Provided that there is a resistance accumulated within very sus-
reasonably wide genetic base, this will ceptible host populations of highly prized
mean that all the polygenes are present in local landraces. No ‘good source’ of resist-
the population, but their frequency is too ance was necessary, and no diminution of
low for much resistance to be expressed in the prized characteristics occurred. When
any of the individuals. The objective of the breeding for horizontal resistance, there-
breeding is to increase these resistance gene fore, the discernible qualities of the genetic
frequencies. resources must be those of yield, quality of
The most resistant plants are selected and crop product and agronomic suitability.
they become the parents of the next screen-
ing generation. Now the most resistant 15.4.4 Population breeding
individuals will possess perhaps 20 percent Recurrent mass selection means that a
of the total polygenes. In the next screening heterogeneous plant population is screened
generation, this percentage is even higher, for the best individuals, which then become
the parents of the next generation. This plants to total plants. This ratio is called
process is repeated some 10–15 times, the selection coefficient. In practice, this
by which time the upper limits of most means that the screening population should
quantitative variables will have been be as large as possible so that perhaps only
reached. In each breeding cycle (i.e. each one plant in a thousand becomes a parent
generation of recurrent mass selection), of the next generation. The possibilities
there should be at least 10 to 20 parents, depend very much on the nature of the
depending on the nature of the crop. These crop. If the plants are small, such as wheat,
parents may be randomly cross-pollinated, rice or beans, it is entirely feasible to use
or hand-pollinated in all combinations, a screening population of some 100 000
again depending on the nature of the crop. plants, but if the population is a tree crop,
Quantitative variables change as a result such as a fruit or nut species, such large
of selection pressures. The term ‘pressure’ populations are not feasible. However, the
is used in the sense of bringing pressure to size of the screening population is not
bear, of coercion or persuasion, and selec- critical, and if a relatively small population
tion pressures can be positive or negative. is necessary because of land or labour
Positive selection pressures lead to restrictions, the breeding programme will
the increase of a variable, while negative require more time, but the deficiency will
selection pressures lead to its decrease. The be no worse than this.
mechanism of these changes is reproductive Should it transpire that the original
fitness. For example, if a heterogeneous genetic base was too narrow to accumulate
host population is susceptible to a parasite, adequate horizontal resistance, new genetic
the most resistant individuals will be material can be added to the screening
parasitized the least and will reproduce population. This may lead to an initial,
the most, while the most susceptible slight loss of horizontal resistance, but the
individuals will be parasitized the most ultimate potential will be improved.
and will reproduce the least. With each A special aspect of quantitative
generation the population as a whole will variables is that they must all be increased
gain resistance as a consequence of this simultaneously. There is little point in
positive selection pressure for resistance. having high levels of horizontal resistance
Conversely, if the parasite is absent from to all of the locally important parasites
the locality in question, as with the maize of except one. Even a single susceptibility
tropical Africa, or because of a functioning will spoil a cultivar, and make spraying
vertical resistance or the use of a pesticide or some other form of artificial control
(i.e. the vertifolia effect; see above), the necessary (see also sub-optimization,
selection pressure for horizontal resistance above). This is a major difference between
will be negative, and the frequency of breeding for single-gene and multiple-gene
genes controlling horizontal resistance characters. Pedigree breeding allows the
will decrease. This happens because any transfer of a single-gene character, such as
unnecessary genetic characteristics tend a resistance, from a wild plant to a cultivar
to decline to a level called the Hardy- by hybridization and back-crossing. A
Weinberg equilibrium. multiple-gene variable cannot be transferred
Positive selection pressure can be in this way because hybridization leads to
increased by increasing the ratio of selected an immediate dilution. Hence the need for
a simultaneous increase of all quantitative approach is feasible include cocoa (Witch’s

variables during population breeding. broom disease caused by Crinipellis
Once the required levels of quantitative pernisciosa) and rubber (South American
variables have been reached, steps can be leaf blight caused by Microcyclus ulei) in the
taken to produce pure lines, clones, or syn- Amazon valley, tea (Blister blight caused
thetic or hybrid varieties, according to the by Exobasidium vexans) in India, date
requirements of the crop in question. palms (Bayoud disease caused by Fusarium
The maize of tropical Africa provided oxysporum f.sp. albedinis) in North Africa,
good examples of both negative and coconut (Cadang-Cadang disease) in the
positive selection pressures to tropical rust. Philippines, white pines (Blister rust caused
It also illustrated the need for horizontal by Cronartium ribicola) in North America,
resistance to all locally important parasites, and many other plantation forest species.
because subsistence farmers do not use
pesticides on their food crops. 15.4.7 Negative screening
Negative screening means that you identify
15.4.5 Male gametocides the worst individuals in a population and
A male gametocide is a chemical that makes remove them, rather than identifying the
a plant male-sterile but female-fertile. By best individuals and keeping them. Negative
using male gametocides in a screening pop- screening can be useful in two situations. The
ulation, inbreeding plants, such as wheat, first occurs when there is a danger of cross-
can be converted to outbreeders. A wheat pollination from undesirable individuals
population then becomes the equivalent of in a screening population, such as open-
a maize population, with unsprayed plants pollinated crops, or self-pollinated crops in
acting as male parents. This can be very which the mother plants have been treated
useful as it eliminates the laborious and with a male gametocide. These undesirable
severely limiting process of cross-pollinat- male individuals must be identified and
ing by hand. Working this way in Brazil, either removed or deflowered.
Beek (1988) obtained millions of wheat A similar situation occurs when a
crosses with only an hour or two of work. heterogeneous population of an open-
The details of male gametocides are pollinated annual crop, such as alfalfa, is
beyond the scope of this chapter and being improved during the process of seed
amateur breeders who decide to use these production. It is often more profitable to
chemicals should get the advice of the remove the relatively few individuals that
specialists who are cooperating in the show the most susceptibility, than it is to
participatory plant breeding. collect the individuals that show the most
resistance. This negative selection should
15.4.6 Screening existing populations obviously be conducted before cross-
With some crops, particularly tree species, pollination becomes possible.
it is possible to find all the resistance we Negative screening can also be used
need by screening existing, heterogeneous profitably in a heterogeneous tree crop.
populations. This was possible with coffee For example, a crop of cocoa might be
in Ethiopia (see below). There is then no heavily diseased with Witch’s broom disease
need for a formal breeding programme as (Crinipellis pernisciosa). A small percentage
such. Other tree crops in which such an of the trees are highly susceptible and
are infecting all the other trees. If the farmer’s crop constituted a natural screen-
most susceptible trees are identified and ing population. Each farmer’s maize then
removed, and all other diseased branches constituted a landrace with exactly the right
are also removed, this parasite interference amount of horizontal resistance to tropical
(see above) will stop and the disease will rust for that latitude and that altitude. This
be controlled. Even if the disease is merely was an example of subconscious selection
reduced in intensity, further negative Because tropical rust is so sensitive to
screenings of the most susceptible trees altitude and latitude, the pathosystems,
will eventually control the disease. This and hence the agro-ecosystems, of tropical
procedure is often far more economical rust are small. However, this extreme of
than a positive screening for resistant trees, environmental sensitivity is unusual, and
followed by a subsequent replanting of the with most crop species the agro-ecosystems
entire crop with these selections. are quite large, and relatively few breeding
programmes are necessary.
15.4.8 On-site selection
The maize of Africa illustrated the 15.4.9 A holistic approach
importance of on-site selection (see The tropical maize in Africa also illustrates
above). Puccinia polysora has maximum the need for a holistic approach. Before the
epidemiological competence at the equator, appearance of tropical rust, the maize had
and at sea level. As latitude increases, the no important pests or diseases. In other
epidemiological competence decreases to words, it had high levels of comprehensive
nothing at sea level at the tropics of Cancer horizontal resistance. That is, it had
and Capricorn. As altitude increases, the adequate levels of horizontal resistance to
epidemiological competence decreases to all the locally important parasites. With the
nothing at the equator at elevations of introduction of this re-encounter parasite,
about 1 200 m. this pathosystem balance was immediately
Maize from the highlands of Kenya, lost, and it required about a dozen
where tropical rust lacks epidemiological generations of selection to restore it.
competence entirely, is extremely It can be argued that pathosystem balance
susceptible when planted at sea level near has been lost in virtually all of our modern
the equator. Conversely, maize in Malawi crops. The objective of participatory plant
was reported to be highly resistant to breeding should be to restore pathosystem
tropical rust but it proved to be very balance in each crop species in each agro-
susceptible when planted near the equator ecosystem. When we consider the many
at sea level. This was environmental erosion different crops and the many different agro-
that occurred because the maize had come ecosystems worldwide, this is too big a task
from an area of minimum epidemiological for professional plant breeders to undertake
competence of the pathogen, where it had on their own, and it is perhaps the best
suffered minimum selection pressure for justification of participatory plant breeding.
resistance. When planted in an area of
maximum epidemiological competence, its 15.4.10 Selection pressures for other
susceptibility was revealed. qualities
Because this maize in tropical Africa was If we were to produce new cultivars that
cultivated as an open-pollinated crop, each had high levels of horizontal resistance to all
locally important parasites, but which had 15.5 EXAMPLES OF BREEDING FOR
reduced yield and quality of crop product, HORIZONTAL RESISTANCE
we would be sub-optimizing (see above). Simmonds (1991) has compiled a compre-
This is why we should use modern but sus- hensive review of the results of breeding for
ceptible cultivars as our genetic resource. It horizontal resistance. He gives examples
is clear that a good source of resistance is of durable resistance in 21 species of crop,
not necessary when breeding for horizontal functioning variously against airborne and
resistance, but that high yield, quality and soil-borne pathogens—fungal, bacterial,
agronomic suitability are necessary. The viral, insect and nematode. Stoner (1992)
levels of various horizontal resistances are reviewed 705 papers on host resistance to
increased while selection pressures for yield insects and mites in vegetables, and she also
and quality are maintained to ensure that quotes reviews of this topic in grain crops,
these qualities are not reduced. alfalfa and cotton. She comments that, in
most studies, the resistance is a quantitative
15.4.11 Measurement of horizontal trait, but she adds that there has been little
resistance plant breeding for resistance to insects.
When measuring the results of breeding for
horizontal resistance, assessments can be 15.5.1 Potatoes in Kenya, Mexico,
relative only. That is, we can say that a new Scotland and the United States of
cultivar has either greater or less horizontal America
resistance to a particular parasite than John S. Niederhauser was one of the
another well known and well tried cultivar. pioneers of horizontal resistance. Indeed,
An alternative description can be given with he was the first scientist to reject the
the phrase ‘spraying not necessary’, but use of vertical resistance in favour of
even this must be qualified with the rider horizontal resistance, and he did this in
that this is only true in a normal season. Mexico with resistance to Late blight of
potato (Phytophthora infestans). His most
15.4.12 Crops that are difficult or famous cultivar was Atzimba, and Mexican
impossible to breed scientists have continued his work. Blight is
For technical reasons, some crops are so severe in Mexico that the popular cultivar
difficult or even impossible to breed, and Alpha has to be sprayed with fungicides 25
amateur breeders should not attempt to times each season. The new, horizontally-
improve them. These include banana, citrus, resistant cultivars, such as Sangema and
date palm, figs, garlic, hops, horseradish, Tollocan, need to be sprayed only once or
olives, pineapple, sisal and wine grapes. twice each season.
However, most of the main food crops Working in Scotland, Simmonds (1976)
are easy to breed, and none of them could demonstrated that the potatoes cultivated in
be described as being difficult to breed. Europe (Solanum tuberosum) were derived
Worldwide, it is clearly logical for amateur from the S. andigena of South America.
plant breeders to work with participatory With only four generations of recurrent
plant breeding of crops that are easy to mass selection he was able produce ‘neo-
breed, while the professional plant breeders tuberosum’ from S. andigena, and he was
should work with crops that are difficult also able to accumulate useful levels of
to breed. horizontal resistance to Late blight.
In Kenya, Robinson (1996) attempted encounter parasite was inadvertently intro-

to imitate the maize of tropical Africa duced. The coffee crops of Ethiopia were
when breeding potatoes for horizontal heterogeneous, and trees with the mini-
resistance to both Late blight (Phytophthora mum horizontal resistance lost all their ber-
infestans) and Bacterial wilt (Pseudomonas ries three months before harvest. Trees with
solanacearum). He was able to have two the maximum horizontal resistance lost
breeding cycles each year with 150 000 no berries at the time of harvest, and they
seedlings in each cycle. When his cultivar occurred with a frequency of about one in a
Kenya Baraka was released to farmers, the thousand. The overall effect was an average
annual potato production of this country yield loss of 40 percent, and this destroyed
increased from less than 10 000 t in 1974, the economic viability of this crop.
to an estimated 1 million tonnes in 2004. About half-a-million trees were exam-
This production was possible without ined and 650 resistant individuals were
any use of crop protection chemicals, and identified. Their first harvests were kept
without any renewal of seed stocks by the for seed, and about 1 000 seedlings were
use of certified seed tubers. It should be germinated from each tree. The first screen-
noted, however, that the temperate viruses ing criterion was for homozygosity, and
of potato lack epidemiological competence only those trees that were ‘breeding true’
in this country, and the Colorado beetle were kept. (Coffea arabica is self-pollinat-
(Leptinotarsa decemlineata) is absent. ing, with about 3 percent of out-crossing).
In the United States of America, Other tests included yield, cup quality and
Fisher, Deahl and Rainforth (2002) have resistance to other pests and diseases. The
been breeding potatoes for horizontal best 25 lines became available as new cul-
resistance to Colorado beetle (Leptinotarsa tivars for farmers only eight years after the
decemlineata) of potatoes and have made programme was initiated (Robinson, 1996).
useful progress after only a few generations
of recurrent mass selection. No single-gene 15.5.3 Beans in Mexico
resistances occur against this insect parasite, Roberto Garcia Espinosa has been using
and this is apparently the first serious recurrent mass selection on black or common
attempt to breed for resistance to it in more beans (Phaseolus vulgaris) in Mexico with
than a century. a view to increasing the levels of horizontal
resistance to all locally important parasites.
15.5.2 Coffee in Ethiopia Commercial crops will yield up to 1 500 kg/ha
Arabica coffee (Coffea arabica) is apparent- if they are properly protected with fungicides
ly an allo-tetraploid that was derived from and insecticides. Beans from the seventh
two diploid species in the area of modern breeding cycle of the recurrent mass selection
Uganda in about 650 CE. It soon died out programme yield 2 400 kg/ha without any
in its centre of origin, but it was taken at use of crop protection chemicals. This work
an early date to Ethiopia, which became has yet to be scientifically described (Roberto
the centre of diversification. A pathogen Garcia Espinosa, pers. comm., 2007).
(Colletotrichum coffeanum) of modern cof-
fee, which causes Coffee berry disease, was 15.5.4 Sugar cane in Hawaii
left behind, and Ethiopia remained free For many years, the sugar cane breeders
of this disease until 1970, when this re- of Hawaii differed from all other cane
breeders in that they used recurrent mass 15.6 ANCIENT CLONES

selection. They produced about three Ancient clones obviously have high levels
million seedlings in each breeding cycle. of horizontal resistance to all their parasites,
These would be reduced to about 600 000 and this resistance has endured for centuries,
on the basis of visual appearances only. even millennia, in crops such as aroids,
The survivors would be tested for sucrose bananas, dates, figs, garlic, ginger, hops,
content, with further massive reductions horseradish, olives, peppercorns, pineapple,
in numbers. As the number of survivors saffron, sisal, turmeric, vanilla, wine grapes
decreased, the complexity of the tests and yams. Some of these clones, such as wine
could be increased. The final result is that grapes, dates and bananas, are now severely
Hawaiian sugar cane yields twice as much as parasitized in some areas, but this is only
any other country, and it does this without because of new-encounter, foreign parasites.
any use of crop protection chemicals, other
than to protect the cut surfaces of the cane 15.7 AUTOCRATIC AND DEMOCRATIC
setts used for planting. PLANT BREEDING
Breeding for vertical resistance is highly
15.5.5 Sweet potatoes in United technical, expensive, difficult, and repeti-
States of America tious. It usually requires a team of scientists
Jones, Dukes and Cuthbert (1976) working in a large institute. Inevitably,
were among the pioneers of horizontal given these problems, fewer cultivars are
resistance breeding when they worked produced than with population breeding,
with sweet potato (Ipomoea batatas) in and it is important that these cultivars have
South Carolina. They used recurrent mass a wide agro-ecological adaptability so that
selection and accumulated good levels of they can be used over as wide an area as
horizontal resistance to several species of possible. Vertical resistance usually has a
insect pests and fungal parasites, as well as very wide adaptability and, coupled with its
improvements in yield and quality. This is complete control of a parasite, this makes it
an easy crop to work with as it possesses an attractive plant breeding approach.
no vertical resistance genes. Vertical resistance breeding was typical
of the green revolution, and the high yield-
15.5.6 Wheat in Brazil ing, ‘miracle’ wheats and rices. While there
Beek (1988) attempted recurrent mass is no question that the increased yields of
selection with wheat in Brazil, using male these cultivars have saved about a billion
gametocides to achieve large numbers human lives, the fact remains that farmers
of random cross-pollinations. He used had little choice of cultivar because there
hydroponics and single-seed descent to were relatively few of these high-yielding
allow late selection. He made good progress cultivars available. A further disadvantage
in accumulating horizontal resistance was that those cultivars that were avail-
to a number of wheat parasites but he able were liable to fail when their vertical
was unable to complete his programme. resistances broke down. Breeding for ver-
Specialists advising amateur breeders in tical resistance also makes farmers totally
a participatory wheat breeding project dependent on the formal seed system.
should regard Beek’s report as essential This approach might be termed
reading. ‘autocratic’ plant breeding, because it is the
breeder, rather than the farmer, who decides REFERENCES

what kind of cultivar is to be bred, and Beek, M.A. 1988. Selection procedures for
which cultivar is to be grown. Its converse durable resistance in wheat. Paper No. 88-2.
is ‘democratic’ plant breeding, in which Agricultural University, Wageningen,
the farmer participates in the breeding, Netherlands. 114 p.
has a choice of resistance type, has a wide Buddenhagen, I.W. 1987. Resistance and
choice of cultivars, and can make their vulnerability to tropical crops in relation
own decisions concerning which cultivars to their evolution and breeding. In P.R.
to grow. Democratic plant breeding is Day, ed. The Genetic Basis of Epidemics
possible with horizontal resistance, which in Agriculture, pp. 309–326. Annals of the
is so easy to use that it can be employed by New York Academy of Science, Vol. 287.
numerous amateur breeders, who cooperate Dent, D. 1988. Insect Pest management.
with professionals in participatory plant Wallingford, UK, CAB International. 604 p.
breeding and plant breeding clubs. Fisher, D.G., Deahl, K.L. & Rainforth, M.V.
2002. Horizontal resistance in Solanum
15.7.1 Plant breeding clubs tuberosum to Colorado Potato Beetle
Plant breeding clubs are made up of amateur (Leptinotarsa decemlineata Say). American
breeders, who might be hobby gardeners, Journal of Potato Research, 79: 281–293.
environmentalists, green activists, farmers, James, W.C., Shih, C.S., Callbeck, L.C. &
students or even schoolchildren. Each club Hodgson, W.A. 1973. Interplot interference
is independent and free to breed any crop it in field experiments with late blight of potato
chooses, for any improvements it chooses (Phytophthora infestans). Phytopathology,
and using any breeding methods it chooses. 63: 1269–1275.
Their primary objective is likely to be the Jones, A., Dukes, P.D. & Cuthbert, F.P. 1976.
production of new cultivars with sufficient Mass selection in sweet potato: breeding for
horizontal resistance to permit cultivation resistance to insects and diseases and horticul-
without any crop protection chemicals, tural characteristics. Journal of the American
and without any reduction in yield, quality Society for Horticulture, 101: 701–704.
or agronomic suitability. This is because Robinson, R.A. 1996. Return to Resistance.
breeding for durable, horizontal resistance agAccess, California, USA. 480 p. Revised
is so easy when compared with breeding 3rd edition (available at http://www.share-
for the ephemeral, but complete, vertical books.ca/eBooks/ReturnToResistance.pdf;
resistance. accessed 11 Feb. 2009).
Plant breeding clubs are particularly Robinson, R.A. 2004. Self-Organising Agro-
useful with participatory plant breeding. A ecosystems. 455 p. (available at http://www.
group of sympathetic crop specialists can sharebooks.ca/eBooks/AgroEcosystems.
cooperate with several plant breeding clubs, pdf; accessed 11 Feb. 2009).
consisting of various categories of amateur Simmonds, N.W. 1976. Neo-tuberosum and
breeders. Possibly the most effective clubs the genetic base in potato breeding. ARC
are university plant breeding clubs made up Research Review, 2: 9–11.
of student-breeders assisted by professors. Simmonds, N.W. 1991. Genetics of horizontal
Greater details have been provided by resistance to diseases of crops. Biological
Robinson (2004). Reviews, 66: 189–241.
Stoner, K.A. 1992. Bibliography of plant resist- Vanderplank, J.E. 1963. Plant Diseases:
ance to arthropods in vegetables, 1977–1991. Epidemics and Control. New York, USA,
Phytoparasitica, 20(2): 125–180. and London, Academic Press. 349 p.
Storey, H.H., Howland (Ryland), A.K., Vanderplank, J.E. 1968. Disease Resistance
Hemingway, J.S., Jameson, J.D., Baldwin, in Plants. New York, USA, and London,
B.J.T., Thorpe, H.C. & Dixon, G.E. 1958. Academic Press. 206 p.
East African work on breeding maize resist- Vanderplank, J.E. 1978. Genetic and Molecular
ant to the tropical American rust Puccinia basis of Plant Pathogenesis. Berlin, Germany,
polysora. Empire Journal of Experimental Springer Verlag. 167 p.
Agriculture, 22: 1–17.
391
CHAPTER 16
Breeding for quantitative variables

Part 3: Breeding for resistance to
abiotic stresses
Stefania Grando and Salvatore Ceccarelli

16.1 INTRODUCTION paradigm shift was needed when selecting

Plant breeding has been very successful for abiotic stresses.
in environments that are either naturally One hypothesis is that cultivars often
favourable or that can be made profitably defined as ‘widely adapted’ are actually
favourable by irrigation and fertilizer and specifically adapted to conditions that are
by chemical control of pests and diseases. at or near the optimum for crop growth.
Cox et al. (1988) estimated that the Therefore the superiority they have in
annual genetic gains in bread wheat in these environments is lost in suboptimal
the United States of America from 1917 environments.
to 1987 have been 16 kg/ha/yr. Russell The objective of this chapter is to discuss
(1984) found that the genetic gain in maize critical problems associated with breeding
hybrids released between 1930 and 1980 for abiotic stresses, to analyse possible
was 54.2 percent. Austin, Ford and Morgan reasons for the limited success breeding
(1989) estimated 38 kg/ha/yr for the genetic has had in stressed environments, and to
gain between 1908 and 1985 in wheat in indicate that participatory plant breeding
the United Kingdom. Similar examples are is one way to overcome the inherent
available in many other crops. difficulties. Most examples are derived from
By contrast, yield improvements have ICARDA’s barley breeding programme for
been very elusive in marginal environments, low-rainfall areas.
to the extent that the role of breeding for
those environments is often questioned. 16.2 MOST COMMON ABIOTIC
What it is not questioned is why it has STRESSES
not been possible to improve agricultural Abiotic stresses are consequences of
production by simply transferring into extremes of physical environment
marginal environments cultivars or comprising climatic stresses, such as
methodologies that have made breeding drought, flood, heat and cold; and soil or
for favourable conditions so successful. As water conditions, such as salinity, metal
a result, the yield of some important staple toxicity and nutrient deficiency. Plants can
crops has shown only modest increases or experience abiotic stresses resulting from
remained virtually unchanged (Ceccarelli the shortage of an essential resource, or
and Grando, 1996; Ceccarelli et al., 2004). from a toxic excess of a substance, or from
This has been attributed to the difficult climatic extremes. In some cases the same
nature of the target environments where resource can impose stress both when in
yields have shown little increase (Passioura, shortage and when in excess (i.e. water and
1986; Blum, 1988) and has been accepted as temperature). Occurrence, severity, timing
inevitable. Therefore, most of the selection and duration of stresses vary from location
work in breeding programmes is done in to location, and in the same location
favourable conditions (Simmonds, 1991), from year to year. In the case of drought,
and much research has been done, and cultivars successful in one dry year may
resources expended, to seek alternatives fail in another, or cultivars resistant to
to empirical breeding for unfavourable terminal drought may not be resistant
conditions, such as analytical breeding and, to intermittent drought, or to drought
more recently, molecular breeding. Much occurring early in the season (Turner, 2002).
less has been done on assessing whether a In addition, abiotic stresses seldom occur in
Breeding for quantitative variables. Part 3: Breeding for resistance to abiotic stresses 393
isolation; they often interact, both with agronomic, genetic, breeding, physiological
other abiotic stresses and with biotic stress. and molecular aspects of drought resistance,
Moreover, areas with a high probability of or as recently more often used, water
abiotic stresses generally have low-input productivity (Passioura, 2006). This is
agriculture (Cooper et al., 1987), because highlighted by the publication of several
the risk of losing the crop or of a low yield reviews (Ceccarelli et al., 2004; Reynolds,
discourages the farmers from using costly Mujeeb-Kazi and Sawkins, 2005; Parry,
inputs, particularly fertilizers. This results Flexas and Medrano, 2005).
in low outputs, poor human nutrition Drought has been always a challenge
and reduced educational and employment to plant breeders, despite many decades of
opportunities, especially for girls. The rural research (Blum, 1993). The development,
poor are particularly badly affected because through breeding, of cultivars with higher
of lack of access to alternative sources of and stable harvestable yield under drought
employment or food. conditions would be a major breakthrough
(Ceccarelli and Grando, 1996). However,
16.2.1 Drought drought resistance is a very elusive trait
Drought, defined as water availability below from a genetic point of view. This is
that required for maximum crop yield, is because the occurrence, severity, timing
one of the main factors limiting crop pro- and duration of drought vary from year
duction. Although it reaches the front pages to year, and although every year there are
of the media as drought warnings or when it “winners”, it is difficult to find those that
causes famine and death, drought is a perma- are consistently successful. To make matters
nent constraint to agricultural production in worse, drought seldom occurs in isolation;
many developing countries, and an occa- it often interacts with other abiotic stresses
sional cause of losses in agricultural pro- (particularly temperature extremes), and
duction in developed ones. Several drought with biotic stress. As mentioned earlier
warnings have been issued in recent years in this chapter, the risk of losing the crop
in Australia, Europe and the United States because of drought limits the use of inputs.
of America. Climate changes will increase Also the definition of dry areas seems
the frequency of droughts, particularly in to be an elusive issue. This is illustrated by
Southeast Asia and Central America, and by the distribution of crops in different agro-
2050 are expected to cause water shortages climatic environments. For example, in a
for 67 percent of the future population in country such as the Syrian Arab Republic,
the world (Ceccarelli et al., 2004). with a large spatial variability of rainfall
In areas where water availability is within short distances (van Oosterom,
limited, and irrigation is not available, the Ceccarelli and Peacock, 1993), bread wheat
choice of crops is restricted to a few, and (Triticum aestivum L.), durum wheat
often to only one, thus making farmers in (T. turgidum var. durum L.) and barley
those areas vulnerable for lack of options. among the cereals, and faba bean (Vicia
In fact, most of the rural poor live in faba L.), chickpea (Cicer arietinum L.) and
areas where crop productivity and crop lentil (Lens culinaris L.) among the food
diversification are limited by lack of water. legumes, are grown in progressively drier
Therefore it is not surprising that there is environments, with some overlapping.
an ongoing global research effort on social, Therefore, a dry area for faba bean or bread
wheat is moderately favourable for durum Soil mineral stresses are increasingly
wheat and chickpea, and a dry area for becoming important limiting factors for
durum wheat and chickpea is moderately crop plants in many parts of the world.
favourable for barley and lentil. Acid soil and associated aluminum toxicity
At the drier end of the spectrum, barley affect over 2 billion hectares worldwide
and lentil are the only rainfed crops, and (Humphreys and Humphreys, 2005).
the other cereals or legumes are only grown Mineral nutrient deficiency can be caused
under supplementary or full irrigation. The by low nutrient status of the soil, low
situation described for the Syrian Arab mobility or availability of nutrients within
Republic applies to most countries of the the soil.
Mediterranean basin and West Asia, and Salinity is generally defined as the
for crops such as millet, sorghum and maize presence of excessive amount of soluble salts
to the dry areas of the tropics, and is only that hinder or affect the normal function of
altered by irrigation. plant growth (Shafiq-ur-Rehman, Harris
The complexity of breeding for dry and Ashraf, 2005). Saline soils have a
areas is not only due to the biological mixture of chloride salts, with sodium
complexity of drought resistance, but chloride being often dominant. Salinity can
also to the consequences of drought for be divided into primary sources in soils
the livelihood of people living in the dry derived from saline parent rocks (Sposito,
areas. In developed countries, farmers have 1989) and secondary salinization caused
various forms of social protection against by human intervention, such as irrigation
the devastating effects of drought, while (Sposito, 1989).
in developing countries farmers have to Salinization is one of the most common
survive on their own, usually selling their forms of soil degradation. Almost all con-
assets, most commonly livestock. In areas tinents have problems related to saline soils
affected by drought in developed countries (Pessarakli, 1999), and is particularly severe
farmers may prefer cultivars capable of in arid and semi-arid regions.
high yields in the few favourable years; It is estimated that 6 percent of the
this is very different in the dry areas of world’s land and 30 percent of the world’s
most developing countries with no or irrigated areas already suffer from salinity
little social assistance where farmers prefer problems (Unesco Water Portal, 2007).
varieties capable of some yields even in the
driest years. This is an example that the 16.2.3 Temperature stresses
same biological problem in different social Temperature extremes can be experienced
contexts requires different solutions. on both a daily or seasonal basis. Long-
term climatic changes lead to higher average
16.2.2 Soil toxicities and deficiencies temperatures and increase the frequency
Soil plays a major role in determining and severity of extreme temperature events.
the amount and availability of nutrients As with other stresses, early and late stages
and toxic minerals. Soil toxicities and of crop growth are particularly sensitive
deficiencies render more than one hundred to temperature extremes. Plants can be
million hectares of agricultural land marginal affected by exposure to prolonged periods
for agriculture, limiting production and of moderately high temperatures as well to
creating poverty for millions. short periods of extremely high tempera-
tures. Low temperatures can affect plants have become more frequent over most land
by chilling, which leads to physiological areas, the frequency of heavy precipitation
and developmental abnormalities, and by events has increased over most areas, and
freezing, which causes cell damage. About since 1975 the incidence of extreme high sea
15 percent of arable land is estimated to be level has increased worldwide.
affected by freezing stress (Dudal, 1976). In conclusion, higher temperatures are
Changes in temperature are the most part of the future climate for which breeders
certain aspect of climate changes. The should breed today.
most recent evidence from the Fourth
Assessment Report on Climate Change 16.3 CHALLENGING CONVENTIONAL
of the Intergovernmental Panel on BREEDING CONCEPTS
Climate Change (IPCC), published in Most plant breeders assume that it is
2007, indicates that the warming of the too slow and too difficult to breed for
climate system is unequivocal, as it is now environments where droughts or other
evident from observations of increases in stresses are unpredictable and variable. The
global average air and ocean temperatures, target is hard to define, and heritability, and
widespread melting of snow and ice, and hence response to selection, is too low to
rising global average sea level. This is achieve meaningful results. Therefore most
shown by (i) 11 of the last 12 years (1995– of the breeding for stress environments has
2006) rank among the twelve warmest been actually conducted using the same
years in the instrumental record of global basic approach that has been very successful
surface temperature (since 1850); (ii) the in areas where lack of water or other abiotic
temperature increase is widespread over stresses is seldom important.
the globe, and is greater at higher northern With few exceptions, most breeding
latitudes; (iii) global average sea level has programmes share the following concepts:
risen since 1961 at an average rate of • selection has to be conducted under the
1.8 mm/yr, and since 1993 at 3.1 mm/ well-managed conditions of research
yr, with contributions from thermal stations. It is felt that environmental
expansion, and melting glaciers, ice caps noises can be kept under control, error
and the polar ice sheets; and (iv) observed variances are smaller and response to
decreases in snow and ice extent are also selection higher;
consistent with warming. Satellite data • cultivars must be genetically homogenous
since 1978 show that annual average Arctic (pure lines, hybrids, clones) and must be
sea ice extent has shrunk by 2.7 percent per widely-adapted over large geographical
decade, with larger decreases in summer of areas;
7.4 percent per decade. Mountain glaciers • locally-adapted landraces must be
and snow cover on average have declined replaced because they are low yielding
in both hemispheres. and disease susceptible;
The 2007 report indicates that it is also • seed of improved cultivars must be
very likely that over the past 50 years, cold disseminated through mechanisms
days, cold nights and frosts have become and institutions such as variety release
less frequent over most land areas, and hot committees, seed certification schemes
days and hot nights have become more and governmental seed production
frequent, and it is likely that heat waves organizations; and
• the end users of new varieties are may be very different from those of
not involved in selection and testing; the breeder (Hardon and de Boef, 1993;
they are only involved at the end of Sperling, Loevinsohn and Ntabomvura,
the consolidated routine (breeding, 1993). Typical examples are crops used
researcher-managed trials, verification as animal feed, such as barley, where
trials), to verify if the choices made for breeders often use grain yield as the
them by others are appropriate or not. sole selection criterion, while farmers are
Breeders have very seldom questioned usually equally concerned with forage
these assumptions. When they have, it has yield and the palatability of both grain
been found that: and straw.
• selection in well-managed research Although the chapter is largely based on
stations tends to produce cultivars that the strategies and methodologies developed
are superior to local landraces only under during the last 20 years in the ICARDA
improved management—not under the barley breeding programme, we believe that
low-input conditions typical of the the main findings have general applicability.
farming systems of stress environments. They will be described to demonstrate
The result is that although many new that it is indeed possible to improve the
varieties outyield local landraces on a production of a typically low-input crop
research station and some are released, such as barley, grown in environments with
few if any are actually grown by farmers low and poorly-distributed rainfall, low
in difficult environments; temperatures in winter, high temperatures
• poor farmers in stress environments tend and drought during grain filling, low soil
to maintain genetic diversity in the form fertility and poor agronomic management.
of different crops, different cultivars The data were mainly obtained from
within the same crop or heterogeneous three locations in the northern Syrian
cultivars, or combinations, to maximize Arab Republic (Tel Hadya, Breda and
adaptation over time (stability), rather than Bouider). They represent three distinct
adaptation over space (Martin and Adams, agricultural systems. Tel Hadya (348 mm
1987a). Diversity and heterogeneity serve average annual rainfall) is a favourable
to disperse or buffer the risk of total crop high-input environment that lends itself to
failure due to environmental variation. a wide choice of different crops. Bouider
This is in sharp contrast to the trend of (236 mm average annual rainfall) represents
modern breeding towards uniformity; the opposite extreme: a typical low-input,
• resource-poor farmers seldom use the high-risk environment where barley is the
formal seed-supply systems. They only possible rainfed crop. Breda (273 mm
frequently rely on their own or on average annual rainfall) is intermediate
neighbours’ seed (Almekinders, Louwaars between the two, located on the edge of
and de Bruijn, 1994). Therefore, when the the area where Arabi Aswad becomes the
appropriate cultivar is selected, adoption dominant landrace. The three sites are
is much faster through non-market geographically close, located 35 (Tel Hadya),
methods of seed distribution (Grisley, 60 (Breda) and 80 km (Bouider) south-
1993); and east of Aleppo. The key aspects of these
• when farmers are involved in the strategies and methodologies are: (i) direct
selection process, their selection criteria selection for specific adaptation in the target
environment (Chapter 9 in this volume); available in many crops, such as lentil

(ii) use of locally-adapted germplasm; (Erskine and Choudhary, 1986), sorghum
(iii) use of plot techniques and experimental (Blum, Golan and Mayer, 1991), bread
design to control environmental variation and durum wheat (Porceddu and Scarascia
(Chapter 3); (iv) participation of farmers in Mugnozza, 1984; Damania and Porceddu,
selection; and (v) reliance on the informal 1983; Spagnoletti-Zeuli, De Pace and
seed-supply system to make the seed of new Porceddu, 1984; Damania, Jackson and
cultivars available to farmers (Chapter 21). Porceddu, 1985; Lagudah, Flood and
Halloran, 1987; Blum et al., 1989; Elings and
16.4 TYPE OF GERMPLASM Nachit, 1991), beans (Martin and Adams,
In breeding for resistance to abiotic stresses 1987a, 1987b), barley (Ceccarelli, Grando
there are certain types of germplasm—lan- and van Leur, 1987; Weltzien, 1988; Asfaw,
draces, wild relatives or wild progenitors— 1989; Weltzien and Fishbeck, 1990) and
that, although of limited or no value in others. Therefore landraces contain a large
breeding for favourable, potentially high- amount of readily-usable genetic variation.
yielding conditions, may play a funda- Selection within landraces is one of the
mental role in the success of a breeding easiest, oldest and cheapest methods of
programme. In many developing countries, plant breeding. In most cases, any interest
landraces (also called farmers’ varieties, shown by researchers in the variability
old cultivars or primitive cultivars) are of landraces has been academic; we know
still the backbone of agricultural systems of few cases in which the variability has
in unfavourable environments (Ceccarelli, actually been used in breeding programmes.
1984; Grando, von Bothmer and Ceccarelli, Yet, as mentioned before, the use of the
2001). In these environments, the replace- variability of landraces in the area where
ment of these cultivars has proved to be landraces are adapted is a cheap and easy
a difficult task. The reasons why farmers way to make progress.
still prefer to grow only landraces or con- At ICARDA we have used a large
tinue to grow landraces even after partial collection of barley landraces made in 1981
adoption of modern cultivars are not well in the Syrian Arab Republic and Jordan
documented, but include quality attributes (Weltzien, 1982). The collection was made
such as food and feed quality, and seed stor- by visiting the fields of 70 farmers and
ability (Brush, 1999). Landraces are often collecting 100 individual heads in each
able to produce some yield even in difficult field. Cvs. Arta, Tadmor and Zanbaka are
conditions, whereas modern varieties are three examples of pure lines identified so
less reliable. For example, where farmers far from two widely grown Syrian barley
have adopted modern cultivars they also landraces (Tables 16.1 and 16.2).
have retained the landraces on the most In developed countries, landraces
unfavourable areas of the farm (Cleveland, have been the basic material for genetic
Soleri and Smith, 2000). improvement in many crops until about
Landraces of self-pollinated species are 50 years ago. In these countries, however,
mixtures of a great number of homozygote the identification of superior genotypes has
genotypes (Brown, 1978, 1979; Ceccarelli probably led many breeders to concentrate
and Grando, 1999; Grando, von Bothmer their attention on those few genotypes and
and Ceccarelli, 2001). Such evidence is this has resulted in: (i) the use of relatively
TABLE 16.1
Grain yield (t/ha) of Tadmor and Zanbaka in 11 and 8 locations respectively in the northern Syrian
Arab Republic
Year Location (Province) Arabi Aswad Tadmor Zanbaka
1991 Shurkrak (Raqqa) 0.220 0.130 -

Al Ayouj (Raqqa) 0.260 0.270 -
Beer Asi (Raqqa) 0.180 0.170 -
1992 Bylounan (Raqqa) 0.640 0.940 1.100
Masadeih (Hassake) 1.350 1.600 1.330
1993 Bylounan (Raqqa) 0.792 1.176 1.132
Shurkrak (Raqqa) 0.666 1.268 0.916
1994 Bylounan (Raqqa) 0.360 0.575 0.530
Al Wastah (Raqqa) 0.560 0.570 0.650
Tell Hamzeh (Hassake) 0.812 0.876 1.250
Al Hamar (Hassake) 1.100 1.000 0.650
Mean 0.631 (0.780) 0.780 0.945
% increase over Arabi Aswad 23.6 22.4
Notes: The mean in parentheses is calculated from the locations in common with Zanbaka. The data are from trials
conducted in farmers’ fields, without fertilizer.
TABLE 16.2
Grain yield (t/ha) of Arta in 51 locations over seven cropping seasons
Year No. of sites cv. Arta cv. Arabi Abiad % increase
1986–1987 1 3.738 2.929 27.6

1988–1989 5 1.814 1.530 18.6
1989–1990 8 2.044 1.747 17.0
1990–1991 11 2.388 2.187 9.2
1991–1992 17 3.336 2.455 35.9
1992–1993 5 2.551 1.958 30.3
1993–1994 4 1.160 1.069 9.4
Note: The data are from trials on farmers’ fields in the Syrian Arab Republic (except those of 1986–1987, which are from
Breda research station).
few parents, leading to a considerable The comparison between barley

reduction of genetic diversity; and (ii) the landraces and modern cultivars under a
loss of most of the landraces before they range of conditions, from severe stress
could be collected and conserved in (low input and low rainfall) to moderately
germplasm banks, and (probably) before favourable conditions (high inputs and
assessing whether their potential had been high rainfall), has consistently indicated
fully exploited. that:
The value of landraces as sources of • landraces yield more than modern
drought tolerance is well documented in the cultivars under low-input and stress
case of barley in the Syrian Arab Republic conditions (Figure 16.1);
(Grando, von Bothmer and Ceccarelli, • the superiority of landraces is not
2001; Comadran et al., 2008; Pswarayi associated only with mechanisms to
et al., 2008) and in several other crops escape drought stress, as shown by their
elsewhere (Brush, 1999). heading date;
FIGURE 16.1
Grain yields (t/ha) of pure lines derived from Syrian landraces and modern cultivars
at three levels of stress in the Northern Syrian Arab Republic
7
6.2
6
5 4.5
4
3.3
t /ha
3.1
3
2
1
1 0.6
P<0.001 n.s. P<0.001

0
Stress Intermediate Non-stress
Level of stress
Modern Landraces
Source: Ceccarelli, 1996.
• within landraces there is considerable rior is based on work conducted in research

variation for grain yield under low-input stations. Even those breeding programmes
and stress conditions, but all the landrace- addressing target environments that have
derived lines yield something, while most low yield potential because of the combi-
modern cultivars fail; nation of biotic and abiotic stresses have
• landraces are responsive to both inputs rarely challenged this assumption.
and rainfall and the yield potential of The superior performance of landraces in
some lines is high, though not as high as dry areas is also evident from the frequency
modern cultivars; and with which they are selected by farmers
• it is possible to find modern cultivars that (Figure 16.2). The data in Figure 16.2 also
under low-input and stress conditions show the importance and the role that
yield almost as well as landraces, but their the wild relatives, in this case the wild
frequency is very low. progenitor of cultivated barley, Hordeum
The data in Figure 16.1 also suggest spontaneum, have as a source of resistance
that selection conducted only in high-input to extreme levels of drought (Grando, von
conditions is likely to miss most of the lines Bothmer and Ceccarelli, 2001; Ceccarelli et
that would have performed well under low- al., 2004).
input conditions. Figure 16.1 also shows Gas exchange observations made
that the assumption of most breeding pro- at anthesis in a wet site showed that
grammes that landraces are genetically infe- H. spontaneum had widely open stomata,
FIGURE 16.2
Frequency of selection by farmers of four types of germplasm (modern, landraces,
landraces × modern and landraces × Hordeum spontaneum)
80
70
60
50
Percentage
40
30
20
10
0
Tel Brack Bylounan J. Aswad B. Sharky
Locations
Modern Landraces Landraces × Modern Landraces × H. spontaneum
TABLE 16.3
Gas exchange parameters of Hordeum spontaneum accessions (means of 12 accessions) and ratio
H. spontaneum/H. vulgare at Tel Hadya, Syrian Arab Republic
Parameter Units H. spontaneum H. spontaneum/H. vulgare
Net photosynthesis μmol m-2 s-1 16.90 ± 0.80 1.49

Leaf conductance mol m-2 s-1 0.40 ± 0.03 3.57
Transpiration efficiency 4.35 ± 0.09 0.63
Leaf temperature Co 25.50 ± 0.36 0.85

Pre-dawn leaf water potential Mpa 0.89 ± 0.25 1.35
higher net photosynthesis and lower pre- only 176 mm rainfall (Grando, von Bothmer
dawn leaf water potential at this stage of and Ceccarelli, 2001).
development than did cultivated barley These lines had some photosynthetic
(Table 16.3). The ability of some accessions activity early in the morning, even though
of H. spontaneum to tolerate extreme levels six times less than in absence of stress,
of drought stress was evident during the stomata were open and the pre-dawn leaf
severe drought of 1987, when two lines of water potential was negative. At the same
H. spontaneum were the only survivors in time, the stomata of the black-seeded local
the breeding nurseries grown at Bouider landrace, Arabi Aswad, considered by
(Syrian Arab Republic), which had received farmers to be very resistant to drought,
were closed, even though the pre-dawn leaf heads are grown as head rows and tested for
water potential was slightly higher than in disease resistance or quality characteristics.
H. spontaneum. By midday, the stomatal Some bulks will lose the superiority shown
conductance of H. spontaneum decreased the year before because of genotype ×
and net photosynthesis became negative, environment interaction and because of
while the stomatal conductance of Arabi decreasing heterozygosity and associated
Aswad was zero. reduced heterotic effects. The corresponding
families will also be discarded.
16.5 BREEDING METHODS The families deriving from the populations
Individual plant selection (such as in the that maintained their superiority for three
widely used ‘pedigree method’) in crops cropping seasons will enter yield testing.
that are normally grown in dense stands When the programme is fully
is very effective for traits that are not implemented, the yield trials contain two
affected by competition. The best example types of materials: new bulks, and pure
is probably disease resistance. However, lines derived from the superior bulks of
many characters of interest to the breeder the previous cycle. If the requirements
are strongly affected by competition. for the genetic uniformity of the varieties
Among others, the ability to tolerate water to be released in a given country are very
stress is certainly greatly affected by the strict, only the pure lines will considered as
distance between plants competing for candidates for release.
limited available water. The result is that The method is based on the basic
isolated plants (such as those of a spaced- assumptions that (i) a superior bulk is
plant F2 used in the pedigree method) grow made by a large number of superior
much better than they would if planted at genotypes, and (ii) that if the superiority is
normal density. The argument that this does maintained for a period of three cropping
not matter as long as all plants are in the seasons in a highly variable environment,
same conditions ignores the possible effects the probability is small that the superiority
of genotype × competition interaction. is associated with heterosis. The method
One breeding method that, in the case can also be used to test the importance
of self-pollinated crops, seems particularly of population buffering in relation to
suitable to breeding for resistance to abiotic stability.
stresses is the bulk-pedigree method, in The method is based on the exploitation
which, after producing the F1 and the F2 on of the genetic variance between
station, three years of multilocation yield populations (Vb) because estimates of Vb
testing and selection of the bulks are carried are comparatively easy and economical to
out in the target environment(s). Selection obtain, while estimates of within-population
is done between bulks by identifying the variance (Vw) are more expensive and
best populations for either yield or other much less precise because of interaction
characters. In parallel with the field testing and competitive effects (Simmonds, pers.
of the bulks, a within-bulks selection is comm.).
conducted only in those bulks that are This method has proved to be ideal for
selected for the next level of field testing: 10 use in participatory breeding programmes
to 50 heads are collected from the selected with self-pollinated crops (Ceccarelli and
populations. The progenies of the selected Grando, 2007).
16.6 SELECTION CRITERIA leaf turgor, abscisic acid content, transpira-

The interest in selecting for traits other tion efficiency, water-use efficiency, carbon
than yield in plant breeding programmes isotope discrimination and re-transloca-
aimed at increasing crop production is tion), and developmental and morphologi-
motivated by the difficulties inherent in cal traits (such as leaf emergence, early
selecting directly for yield. Literature on growth vigour, leaf area index, leaf waxi-
the inheritance of yield in several crops has ness, stomatal density, tiller development,
led to the conclusion that yield is inherited flowering time, maturity rate, vernalization
in a complex manner (Blum, 1988). In spite requirement and root characteristics).
of the widespread reference to ‘yield genes’, In the case of barley, traits more
it is evident that yield as such is not under consistently associated with higher grain
‘direct genetic control’. Rather, it is the yield under drought are growth habit, early
multitude of physiological and biochemical growth vigour, earliness, plant height under
processes—the integrated effect of which is drought, long peduncle and a short grain-
measured as yield—that are under genetic filling duration (Acevedo and Ceccarelli,
control (Blum, 1988). The complex manner 1989). Three traits that deserve a special
of inheritance of yield is evident from the mention are leaf epidermal conductance
generally low estimates of heritability that (Sinclair, 2000), osmotic adjustment
are of common occurrence for characters (Serraj and Sinclair, 2002) and desiccation
that are under complex genetic control. The tolerance (Ramanjulu and Bartels, 2002),
difficulties of selecting for yield become even which appear related to survival under
greater in environments characterized by severe stress conditions. Even though the
unpredictable variability in the frequency, low yields resulting from survival traits
timing and severity of a number of climatic may look irrelevant from the perspective
stresses. of high-input agriculture, they are crucial
Breeding for drought resistance based to the livelihood of farmers in some of the
on putative traits (defined as traits associ- driest regions of the world.
ated with drought resistance, but easier to While the analytical approach has been
select for than grain yield) has been, and very useful in understanding which traits are
still is, very popular (Richards et al., 2002). associated with drought tolerance and why,
The ideal trait to be used as an additional it has been less useful in actually developing
or alternative selection criterion to yield in new cultivars showing improved drought
breeding for stress conditions should satisfy resistance under field conditions. Under
the following requirements: (i) be causally such conditions, drought varies in timing,
related or genetically linked to yield under intensity and duration, and therefore it is
stress conditions; (ii) exhibit genetic vari- the interaction among traits to determine
ation; (iii) be highly heritable; and (iv) be the overall crop response to the variable
easy, inexpensive and quick to screen for. nature of the drought stress rather than the
Traits that have been investigated include expression of any specific trait (Ceccarelli,
physiological and biochemical traits (such Acevedo and Grando, 1991). A typical
as osmotic adjustment, proline content, sto- example is offered by early growth vigour,
matal conductance, epidermal conductance, a trait that is unanimously considered
cell membrane stability, cell wall rheology, important in reducing the amount of
canopy temperature, relative water content, water lost by evaporation from the soil
surface, and therefore in increasing water- While there is currently substantial

use efficiency (Richards et al., 2002) in investment in molecular approaches to the
crops grown on current rainfall (absence study of drought resistance, there are not yet
of stored moisture). The study of barley success stories based on the identification
landraces from the Syrian Arab Republic of specific genes and their utilization for
has revealed that genotypes with a modest this challenge (Chapman, 2008).
early vigour can successfully achieve the Ultimately it is the drought resistance
same result with a prostrate growth habit under field conditions that needs to be
(Ceccarelli and Grando, 1999). improved. Yield under stress conditions
Furthermore, many of the studies continues to be the major selection
on putative traits have been conducted criterion. In the case of barley, additional
independently from, or as a side-activity selection criteria utilized are early growth
to, breeding programmes, and by non- vigour, plant height under stress, tillering
breeders. As a consequence, in general, and earliness.
breeders have taken a sceptical attitude
towards these studies, with the well- 16.7 ARCHITECTURE OF GENOTYPES
founded justification that the stated AND YIELD STABILITY
conclusions are affected by either the low One of the most dramatic changes
number of genotypes involved, or the introduced by modern agriculture has been
particular type of germplasm used or the reduction of variability. The narrowing
insufficient number of environments. This of the genetic base that has been a feature
attitude emerges clearly even in the case of of plant breeding in developed countries
individual-trait breeding to enhance genetic has been accompanied by a trend towards
yield potential (Rasmusson, 1987). homogeneity: one clone, one pure line, one
Breeding for drought resistance based hybrid (Simmonds, 1983). Uniformity and
on direct selection for grain yield in the broad adaptation are very useful attributes
target environment (empirical or pragmatic to accommodate large-scale centralized
breeding) appears intuitively to be the most seed production (Davis, 1990). While this
obvious solution. However, it has faced trend is now being questioned in developed
the major criticism that since field-drought countries (Wolfe, 1992), it is still very
is such a moving target, the chances of common in breeding programmes for
progress appear slow at best and possibly developing countries at both national and
remote. One major consequence of this international levels.
attitude has been to study a less mobile In breeding programmes aiming at
target by simulating drought in laboratory increased stability, the problem of reduced
(or greenhouse) conditions, which results variability is particularly serious in relation
in generally irrelevant shocks (Passioura, to the two major genetic mechanisms
2002). Several studies have been and are promoting stability: individual buffering and
being conducted addressing ‘laboratory population buffering. Individual buffering
drought’ with the main justification is largely a property of heterozygotes,
being to discover mechanisms and genes and although there is some evidence of
(Yamaguchi-Shinozaki and Shinozaki 1994; individual buffering not associated with
Kasuga et al., 1999; Nakashima et al., 2000; heterozygosity, it may be difficult to
Seki et al., 2001). exploit this mechanism in self-pollinated
diploid crops (Allard and Bradshaw, 1964). that, during millennia of cultivation under
However, as modern varieties of cereal adverse conditions, natural and artificial
crops such as wheat and barley are mostly selection have not been able to identify
pure lines, they must rely on individual either an individual genotype possessing
buffering to be stable. Population buffering a key trait associated with its superior
is a mechanism of stability associated with performance, or an individual genotype
genetic heterogeneity. ‘Varieties’ made with a specific architecture of different
up of a number of genotypes, such as traits. On the contrary, the combined
the landraces, are well buffered (stable), effects of natural and artificial selection
because each member of the population is has led to diversity in architecture of
best adapted to slightly different conditions genotypes, representing different
from other members of the population. The combinations of traits. These populations
stability of the individuals is sacrificed to can be extremely useful for understanding
maximize the stability of the population. mechanisms that enhance stability in stress
Although a direct relationship between environments, not only from the genetic
genetic heterogeneity and stability has yet structure point of view, but also for
to be demonstrated for landraces, it can understanding the adaptive role of given
be speculated that, being the product of traits. In fact, although variable, landraces
natural and artificial selection following grown in environments characterized by
domestication, the genetic structure of a high frequency of stress conditions tend
landraces must bear some advantage, or at to present a high frequency of a given
least cannot be a purely random outcome. expression of specific traits.
The genetic structure of landraces, For example, barley lines extracted from
therefore, may be considered an landraces collected from five sites in the
evolutionary approach to survival and Syrian steppe (Table 16.4) were compared
performance under arid and semi-arid with barley lines extracted from landraces
conditions (Schulze, 1988). It follows collected in Jordan and with a wide range of
TABLE 16.4
Mean of morphological and developmental traits in 1041 modern barley genotypes (unrelated to
Syrian or Jordanian landraces) compared with 322 pure lines extracted from Syrian landraces and
232 pure lines from Jordanian landraces
Traits Modern (n=1041) Landraces
Syrian Arab Republic
Jordan (n=232)
(n=322)
1. Early growth vigour 2.5 b 3.2 a 2.4 b
2. Growth habit 2.8 c 4.0 a 3.1 b
3. Cold tolerance 3.0 a 1.3 c 2.3 b
4. Days to heading 117.9 b 121.2 a 116.9 c
5. Grain filling 39.3 a 35.5 c 37.4 b
6. YP (t/ha) 4.398 a 3.293 c 3.947 b
7. YD (t/ha) 0.483 c 0.984 a 0.835 b
Notes: (i) Traits 1, 2, 4, 5 & 6 were scored or measured at Tel Hadya in 1987/88 (504.2 mm rainfall); trait 3 was scored at
Bouider in 1987/88 (385.7 mm rainfall); and trait 7 was measured at Bouider in 1988/89 (189 mm rainfall), on 521 modern
lines, 92 Syrian landraces, and 86 Jordanian landraces. Early growth vigour (1=good; 5=poor), Growth habit (1=erect;
5=prostrate), Days to heading (days from emergence to awn appearance), Grain filling duration (days between heading and
maturity), YP = Yield Potential, YD = Yield under Drought.
(ii) Means followed by the same letter are not significantly (P<0.05) different based on t-test for samples of unequal size.
modern (non-landrace) barley genotypes. each of these traits. The variability around a
The Syrian lines showed a higher frequency mean expression of each character—which
of genotypes with prostrate or semi- already allows a high degree of adaptation—
prostrate growth habit, cold tolerance might perhaps be considered as a fine-tuning
and short grain-filling period, and a lower mechanism to cope with environmental
frequency of genotypes with good growth fluctuations. Thus, 321 lines derived from
vigour and early heading. Their average Syrian landraces were classified according
grain yield in unfavourable conditions (at to the score for early growth vigour in
Bouider in 1989) was 0.984 t/ha (ranging three classes: good vigour (score <2.5);
from 0.581 to 1.394 t/ha), more than twice intermediate (score = 2.5–3.5); and poor
the average grain yield of modern genotypes vigour (score >3.5). Each class was then
(0.483 t/ha, ranging from crop failure to classified according to the score for growth
1.193 t/ha). The average yield in favourable habit (erect <2.5; semi-prostrate = 2.5–3.5;
conditions of the Syrian landraces (3.293 t/ prostrate >3.5). No genotypes were found
ha) was 75 percent of the average yield in the good vigour-erect, intermediate
in favourable conditions of the modern vigour-erect, poor vigour-erect, and poor
germplasm (4.398 t/ha). vigour-semi-prostrate classes (Table 16.5).
Although this particular set of data is The groups were compared not only for
based on one environment only, it confirms the two traits used in their classification,
the existence of the trade-off between yield but also for days to heading, cold tolerance
in unfavourable conditions and yield in and length of the grain-filling period. Lines
favourable conditions discussed earlier. with good early growth vigour tend to be
Landraces collected in Jordan, from sites less cold tolerant, earlier and with a longer
with milder winters than the Syrian steppe, grain-filling period. This small percentage
have a higher frequency of genotypes that of genotypes will presumably have a yield
have better early growth vigour, more erect advantage in years with slightly milder
habit, less cold tolerance, slightly longer winter temperatures, absence of late frosts
grain-filling period and earlier heading and less severe terminal stress. The highest
than Syrian landraces. Their average frequency of genotypes (71.3 percent)
grain yield in unfavourable conditions combines intermediate early-growth vigour
was only slightly lower (0.835 t/ha) than with semi-prostrate or prostrate growth
Syrian landraces, while their average yield habit. These genotypes are slightly more
in favourable conditions (3.947 t/ha) was cold tolerant than the first group, but
in between the Syrian landraces and the are slightly later in heading. However,
modern germplasm. Syrian landraces they are better equipped to escape terminal
therefore show a combination of escape drought because of the shorter grain-
(early maturity) and avoidance (prostrate filling period. About one-quarter of the
habit and cold tolerance result in good genotypes (22.1 percent) have poor early
ground cover) mechanisms. growth vigour but a very prostrate growth
In addition to the high frequency of habit (growth habit score = 4.2) and a high
combinations of escape and avoidance level of cold tolerance (1.3). Their slightly,
traits, landraces possess another powerful although significantly, later heading is not
mechanism. They are composed of a number necessarily a negative attribute, mostly
of genotypes with a variable expression for because it is compensated for by a very
TABLE 16.5
Frequency of different combinations of early growth vigor (GV), and growth habit (GH), and mean
values of cold tolerance (CT), days to heading (DH) and length of the grain filling period (GF) in a
sample of 322 lines of barley collected in the dry areas of the Syrian Arab Republic (from same trials
as indicated in notes of Table 16.4)
Groups % GV GH CT DH GF
1. Good vigour–Erect 0.0 - - - - -

2. Good vigour–Semiprostrate 1.2 2.2 3.3 1.6 118.8 37.4
3. Good vigour–Prostrate 5.3 2.4 3.9 1.4 119.8 36.6
4. Intermediate vigour–Erect 0.0 - - - - -
5. Intermediate vigour–Semiprostrate 6.2 2.9 3.4 1.5 119.7 35.8
6. Intermediate vigour–Prostrate 65.1 3.1 4.0 1.4 121.2 35.4
7. Poor vigour–Erect 0.0 - - - - -
8. Poor vigour–Semiprostrate 22.1 3.9 4.2 1.3 121.9 35.4
9. Poor vigour–Prostrate 22.1 3.9 4.2 1.3 121.9 35.4
Least Signifigant Difference – LSD0.05 0.2 0.1 0.1 0.6 0.7
Notes: All collection sites are included in the Palmyra region, as defined by Weltzien (1988).
short grain-filling period. In an environment • it is the interaction among these, and

characterized by a combination of different possibly other traits, that plays a key role
abiotic stresses with varying intensity and in determining the differences in overall
frequency every year, a population with performance rather than the expression
such architecture of genotypes is probably of any one of them taken in isolation;
the best solution to long-term stability • because of the interactions among traits,
(Ceccarelli, Acevedo and Grando, 1991). different combinations of traits are
The evidence that, at least in the short expected to produce the same effect in
term, some individual genotypes (pure terms of final yield;
lines) are able to show the same degree of • the role of each individual trait, even
stability as local heterogeneous populations within the restricted terms of reference
has been presented earlier (the examples that have been chosen, depends on the
of cvs. Tadmor and Arta). Even so, the frequency, timing and severity of stresses,
use of population buffering in addition to and on the type of stress; therefore, efforts
individual buffering offers scope for further to identify individual traits causally
increased stability. associated with yield stability under
In conclusion, the evidence discussed stress are unlikely to be successful;
suggests that: • in this type of stress environment,
• genetic differences in yield and yield ‘drought resistance’, defined in terms of
stability under conditions of low winter yield under stress, is a genetic abstraction
temperatures and moisture stress are as much as yield in general;
associated with differences, among others, • analytical breeding to enhance yield
in morphological and developmental stability in stress environments has to
traits such as growth habit, cold tolerance, consider individual traits as part of an
growth vigour and time to flowering. In architecture, rather than in isolation;
other types of stress environments and/ and
or in other crops the suite of traits will • long-term and sustainable improvements
obviously be different; of yield stability should probably
be based on population buffering as not exclude breeding material with wide

achievable with mixtures of genotypes spatial adaptation, while selection for wide
representing different, but equally adaptation tends to eliminate breeding
successful, combinations of traits, as material with specific adaptation.
occurs in landraces. Decentralized selection is different
from decentralized testing, which is a
16.8 PLOT TECHNIQUES AND common feature of breeding programmes
EXPERIMENTAL DESIGNS and takes place, usually in the form of
When genotypes are compared at increasing multilocation trials and on-farm trials, after
levels of moisture stress, small variations in a number of cycles of selection in one or
soil depth, texture and topography have few environments (usually with high levels
increasingly large effects on plot-to-plot of inputs).
variability because of associated differences Decentralized breeding is a powerful
in soil moisture availability. Therefore, it means to adapt crops to the physical
becomes essential to adopt plot techniques environment. However, to exploit fully
and experimental designs that can minimize the potential gains from specific adaptation
these effects. to low-input conditions, breeding must
Various plot techniques to increase the be decentralized from research stations to
efficiency of direct selection in the presence farmers’ fields. Although decentralization
of abiotic stresses have been discussed and farmer participation are unrelated
extensively in Chapter 3 of this volume. concepts, decentralization to farmers’ fields
almost inevitably leads to the participation
16.9 DECENTRALIZED-PARTICIPATORY of farmers in the selection process (Ceccarelli
SELECTION and Grando, 2002).
The term ‘decentralized selection’ was first
used by Simmonds (1984) and defined 16.10 DECENTRALIZED-PARTICIPATORY
as selection in the target environment(s). PLANT BREEDING
Decentralized selection becomes selection The implementation of decentralized-
for specific adaptation when the selection participatory plant breeding (PPB)
criterion is the performance in specific programmes started in 1997 with the
environments rather than the mean aim of developing an alternative way of
performance across environments. Selection conducting plant breeding that is much
for mean performance across a number of more efficient and much quicker in bringing
environments (years and locations) tends to new varieties to farmers, and ensures that
exclude breeding material that performs very the new varieties are adapted to farmers’
well in the lowest yielding years or locations specific environments and end-uses.
but not particularly well in the highest The emphasis of the programme
yielding years or locations, unless data are has been on dry areas, even though the
standardized. On the contrary, selection for approach can also be beneficial to high-
the highest yielding breeding material in rainfall environments.
specific locations or areas will automatically The programme, which has been
include breeding material performing described in detail by Ceccarelli and Grando
well across all locations. In other words, (2007) and by Ceccarelli, Grando and Baum,
selection for specific spatial adaptation will (2007), is based on the following concepts:
• the trials are grown in farmers’ fields using is gained as part of the selection process.
the host farmer’s agronomic practices; Last, but not least, the public investment in
• selection is conducted by farmers in seed production is nearly always paid off
farmers’ fields, so that farmers are the by farmers’ adoption.
key decision-makers; and The programme started in the Syrian
• the traditional linear sequence of Scientist Arab Republic in 1996 and was expanded to
→ Extension → Farmers is replaced by a Algeria, Egypt, Eritrea, Islamic Republic of
team approach, with Scientists, Extension Iran, Jordan, Morocco, Tunisia and Yemen,
Staff and Farmers participating in all using the same bulk-pedigree method
major steps of variety development (see described earlier. Four types of impact can
Figure 9.1). be observed, considered below.
In a conventional breeding programme,
the most promising lines are released as Variety development
varieties, their seed is produced under New varieties were spontaneously
controlled conditions (certified seed) and disseminated from farmer to farmer as
only then can farmers decide whether to early as three years after starting the
adopt them or not. In many developing programme. In the Syrian Arab Republic,
countries the process results in many several thousand hectares are planted with
varieties being released but only a small two varieties, and 12 varieties have been
fraction being adopted. The major adopted by farmers and are under seed
consequence of the PPB concept is that multiplication (Table 16.6). Varieties are
the process transforms the delivery phase adopted both in dry areas and in wetter areas
of a plant breeding programme from being in a much shorter time than in a conventional
supply driven to being demand driven. breeding programme. It also confirms the
Under PPB, it is the initial farmers’ importance of landraces (Tadmor, Arta,
preference that drives the decision of which SLB and JLB lines, Zanbaka, A. Abiad
variety to release. As a consequence, adoption and A. Aswad) as well as H. spontaneum
rates are higher, and risks are minimized, as when farmers’ opinion becomes part of the
intimate knowledge of varietal performance breeding process.
TABLE 16.6
Varieties adopted from the PPB programme by farmers in the Syrian Arab Republic
Pedigree Name Location Rainfall
H.spont.41-1/Tadmor Raqqa-1 Bylounan 212.4

Arta//H.spont.41-5/Tadmor Raqqa-2 Bylounan 212.4
Zanbaka/JLB37-064 Karim Bylounan 212.4
Tadmor/3/Moroc9-75/ArabiAswad//H.spont.41-4 Akram Bylounan 212.4
Mo.B1337/WI2291//Moroc9-75/3/SLB31-24 Suran-1 Suran 383.7
ChiCm/An57//Albert/3/Alger/Ceres.362-1-1/4/Arta Suran-2 Suran 383.7
ER/Apm//Lignee131/3/Lignee131/ArabiAbiad/4/Arta Suran-3 Suran 383.7
Hml-02/5/..Alger/Ceres362-1-1/4/Hml Nawair-1 Suran 383.7
Hml-02/5/..Giza 134-2L/6/Tadmor Nawair-2 Suran 383.7
SLB03-10/Zanbaka Yazem J. Aswad 226.4
Tadmor//Roho/Mazurka/3/Tadmor Salam J. Aswad 226.4
ArabiAswad/WI2269/3/ArabiAbiad/WI2291//Tadmor /4/Akrash//WI2291/WI2269 Ethiad J. Aswad 226.4
Note. Rainfall is annual rainfall in mm, average of the period 2000–2005.

Institutional even within the same location (as shown in

In several countries, PPB has generated Table 16.6) in response to different envi-
considerable change in the attitude of poli- ronmental constraints and users’ needs.
cy-makers and scientists towards the bene-
fits of participatory research, and generated 16.11 DROUGHT-RESISTANT LINES
changes in national breeding programmes. PPB was not specifically designed to breed
for drought tolerance, but rather to adapt
Farmers’ skills and empowerment the crops to a number of environmental
The cyclic nature of the PPB programmes and agronomic conditions and to farmer
has considerably enriched farmers’ preferences. These also include situations
knowledge, improved their negotiation where drought stress occurs frequently
capability, and enhanced their self esteem. and can be very severe. Therefore it is not
By the same token, scientists (breeders) have surprising that the PPB programme has
been enriched by the farmers’ indigenous produced the breeding material with the
knowledge of the crops they grow and the highest level of drought tolerance.
environments in which they grow them. In this section we will give two examples
of lines specifically adapted to dry areas.
Enhancement of biodiversity The first example refers to lines selected in
Different varieties have been selected in the Syrian Arab Republic in 2000, when the
different areas within the same country, and total rainfall in most areas of the country was
FIGURE 16.3
Grain yield (t/ha) of two lines with improved drought resistance in comparison
with the local landrace tested in 2004, 2005, and in 2006 in rainfed location in
the Syrian Arab Republic receiving less than 200 mm of total rainfall
1.2
0.8
t/ha
0.6
0.4
0.2
0
2004-1 2004-2 2005 2006-1 2006-2 mean 2004-1 2004-2 2005 2006-1 2006-2 mean
Improved Local
Line 1=H.spont.41-1/Tadmor//SLB45-090/H.spont.41-2/3/H.spont.41-1/Tadmor//H.spont.41-1/Tadmor
Line 2=Arta/3/Arar/H.spont.19-15//Hml
below average and crop yields were severely and between 0.5 and 3.0 t/ha of biomass
affected. In some areas, the rainfall was so yield (Ceccarelli et al., 2004).
low that the crop did not even germinate; in Two of these lines were tested by farmers
many others the crop failed to produce grain. on large areas (5–20 ha) in 2004, 2005 and
The PPB trials, planted in eight farmers’ 2006, which were all very dry (Figure 16.3).
fields in the Syrian Arab Republic, were In comparison with the local landrace,
affected by different intensities of drought. which itself is considered to be drought
At one extreme, the rainfall was only 50 mm resistant, the two lines showed an average
in the entire season and no germination yield advantage of 44 percent, ranging from
occurred. At the other extreme, the rainfall 9 percent to 67 percent. This includes one
was 252 mm rainfall and average grain yield case in which the local landrace failed, and
was 1.8 t/ha (ranging from 1.0 to 3.2 t/ha). one of the improved lines yielded 0.5 t/
The driest sites, where some new barley ha. Both lines are derived from crosses
entries were able to produce some grain with a pure line of H. spontaneum (the
or some biomass, received between 87 and wild progenitor of cultivated barley), an
130 mm. Average grain and biomass yield indication that some H. spontaneum lines
were very low but some lines were able to can contribute significantly to enhance the
produce between 0.3 and 0.5 t/ha of grain drought resistance of cultivated barley.
TABLE 16.7
Rainfall (mm) and average yield (in parenthesis in t/ha) in five locations in the dry areas of the
Syrian Arab Republic during the three years 2003–2005
Location No. of lines 2003 2004 2005 Mean annual
rainfall
2000–2005 (mm)
Bylounan 8 187 (1.249) 217.4 (0.804) 196 (0.604) 212.4

J. Aswad 6 215 (1.152) 245.3 (0.808) 238 (0.853) 226.4
Melabya 10 275.9 (1.496) 176 (0.432) 187.5 (0.466) 186.5
Siebatt 12 308 (1.406) 319 (0.478) 263 (1.027) 248.8
Al Bab 10 408 (1.355) 296 (0.422) 289.5 (1.084) 307.5
TABLE 16.8
Grain yield (as percentage of the local check) of the highest yielding lines during 2003, 2004 and
2005 in five dry locations in the northern Syrian Arab Republic receiving between 186 and 308 mm
total annual rainfall
Location Line 2003 2004 2005 Mean
Bylounan Arta/SLB22-74 1.109 1.030 1.065 1.068

Bylounan ArabiAbiad/Arar//H.spont.41Arta/….. 1.087 1.100 0.993 1.060
Bylounan Arta//H.spont.41-5/Tadmor/3/SLB05-096 1.098 1.102 0.982 1.060
J. Aswad SLB28-53/SLB21-81 1.030 1.121 0.967 1.039
Melabya Roho/4/Zanbaka/3/ER/Apm//Lignee131 1.151 1.045 1.071 1.089
Melabya ArabiAbiad/Arar//H.spont.41Arta/….. 0.957 1.038 1.101 1.032
Siebatt SLB21-81/SLB22-74 1.216 1.085 1.023 1.108
Siebatt Anadolu86/Sara-02//Zanbaka 1.186 0.993 1.128 1.102
Siebatt Zanbaka/SLB21-81 1.180 1.038 1.036 1.084
Siebatt Sara-01/Sara 1.433 1.009 0.998 1.147
Al Bab ChiCm/An57//Albert/3/Alger/Ceres.362 1.323 1.535 1.207 1.355
Al Bab SLB28-53/SLB21-81 1.282 1.472 1.300 1.352
The second example derives from trials first and Al Bab and J. Aswad the second).
conducted in dry locations in the northern As these lines are the product of one cycle
Syrian Arab Republic during the period of selection, further progress is expected
2003–2005. Two locations (Bylounan and with additional cycles of recombination
Melabya) represent some of the driest areas and selection.
in the Syrian Arab Republic, where barley
is the only possible rainfed crop; J. Aswad 16.12 CONCLUDING REMARKS
and Siebatt are in slightly wetter areas, The objective of this chapter has been to
while Al Bab is a location characterized by discuss what plant breeders can do when
colder winters than the other four. In the the target environment of their breeding
two driest locations, rainfall varied from programme is characterized by chronic
176 mm to 245.3 mm total annual rainfall low yields due to numerous factors,
and average grain yield from 0.432 to 1.496 t/ such as climatic, nutritional and abiotic
ha during the testing period. In the two stresses. The data are mostly derived from
wetter locations, rainfall varied from 215 barley and from one type of dry area (dry
to 319 mm total annual rainfall and average Mediterranean with cold winters and hot
grain yield from 0.478 to 1.406 t/ha, while summers, and crops grown on current
Al Bab was the wettest of the five locations rainfall). However, the paper illustrates
but not the highest yielding because of the some general concepts that, with some
low temperatures in winter (Table 16.7). modifications, could be useful in other
In the five locations, we tested between crops and in other types of dry area.
6 and 12 lines (including the checks) repre- The first concept is that in these
senting the result of two cycles of decentral- environments, climatic, nutritional and
ized participatory selection starting from a biotic stresses usually occur together
common set of 165 lines. The yield of the (though not necessarily all of them all of the
best lines is shown in Table 16.8, expressed time); and, so far, there is little substitute
as a percentage of the local check. for actually exposing the breeding material
At the two driest sites, Bylounan and to a real field situation. Although little
Melabya, five lines outyielded the local practiced, the idea is not new. Nearly
check on average over 3 years by between forty years ago Hurd (1971) published
3.2 percent and nearly 9 percent, but only a paper with the title: Can We Breed for
three lines were consistently superior to the Drought Resistance? The first sentence of
local check in each of the three years. In the the paper was “My answer to the above
two wetter locations, five lines outyielded very pertinent question is a confident and
the local check by between 7.5 percent optimistic ‘Yes’”. He concluded: “One
and 14.7 percent, but only the two lines method is to grow large populations in
in J. Aswad and two of the four lines in early generations under typical dry growing
Siebatt consistently outyielded the local conditions.” Twenty years later, Bramel-
check. In Al Bab, two lines consistently Cox et al., (1991) recognized that the key
outyielded the local check by slightly more to increased production with fewer external
than 35 percent; two lines (ArabiAbiad/ inputs would be through a re-evaluation of
Arar//H.spont.41 and SLB28-53/SLB21- the identification and use of selection and
81) were among the highest yielding lines in testing environments.
two locations (Bylounan and Melabya the Although this concept is obvious to
many and not new, selection for stress stress environment, it is probably for that
environments is still seldom done in the reason.
target environments and it still a highly The main conclusion of this paper is
controversial issue, as it is the relationship that breeding for stress environments is
between high yield under optimum possible, provided it is conducted with
conditions and high yield under abiotic strategies and methodologies that little have
stress conditions (see, for example, Chapter in common with those used in breeding
18 in this volume). This may not always for favourable environments. Adaptation
be necessarily a deliberate choice of one over time can be improved by breeding
breeding strategy or another, but is simply for specific adaptation to a given type of
due to the distance of suitable selection sites stress environment. This can be achieved by
from main cities, with all the associated taking advantage of the temporal variability
inconveniences. We hypothesize that in of stress environments, which permits
these cases an interesting solution may exposure of the same breeding material to
be offered by farmers’ participation in variable combinations of stresses over a
breeding (Ceccarelli and Grando, 1997; (relatively) short period and to accumulate
Ceccarelli, Grando and Baum, 2007). favourable alleles at the several loci involved
Conducting selection in farmer’s fields in drought resistance through successive
has the advantage of exploiting genetic cycles of recombination and selection.
differences under farm conditions, with the We are aware that this is fundamentally
additional advantage of making use of the different from the modern trend of plant
farmer’s knowledge of the crop. breeding towards broad adaptation over
The second concept is the use of space. The difference represents the
germplasm usually ignored by most plant contrasting interests of farmers and seed
breeders, such as landraces and wild relatives. companies. Farmers are interested in
This approach is a direct consequence of cultivars that are consistently superior on
choosing to work in the target environment their farm, regardless of how they perform
and has led to the development of a number at other locations or in other countries.
of barley cultivars, now grown in a number Seed companies, however, want to market
of farmers’ fields in the central and northern as much seed of as few cultivars as possible.
Syrian Arab Republic and in environments Breeders have been breeding, perhaps
considered too difficult and therefore unconsciously, more for seed companies and
beyond the plant breeder’s domain. for their personal prestige than for farmers.
The third concept is that in dry areas, The two objectives coincide when selection
every effort should be made to control and target environments are similar, but
environmental variability in trial and this approach has by-passed millions of
nurseries evaluation. When working at small farmers in difficult environments.
stress sites, the breeder should forget the Recent advances in plant genomics have
typical research-station style of work. The enabled one to dissect various molecular
methodology, experiment designs and plot mechanisms (signal transduction pathways)
techniques used in the very homogeneous involved in drought, cold and salt stress
environment of the experiment station are tolerance and in identifying various
not suitable; in fact, when the conclusion is genes involved in such stress tolerance.
reached that progress cannot be made in a Information generated in genomics should
be integrated into practical plant breeding. Austin, R.B., Ford, M.A. & Morgan, C.L. 1989.
Various genes identified, in both model Genetic improvement in the yield of win-
plants and crop plants, could be used in ter wheat: a further evaluation. Journal of
future for developing stress-tolerant plants Agricultural Science, Cambridge, 112: 295–
through either marker-assisted selection or 301.
direct gene transfer. Blum, A. 1988. Plant breeding for stress envi-
ronments. Boca Raton, USA, CRC Press.
ACKNOWLEDGEMENTS Blum, A. 1993. Selection for sustained pro-
This chapter is based on the work carried duction in water-deficit environments. In
out at ICARDA over more than twenty International Crop Science I, pp. 343–347.
years. Many people have made this work Madison, USA, CSSA.
possible, and we would like to acknowledge Blum, A., Golan, G., Mayer, J., Sinmena, B.,
the technical assistance of Mr A. Ayyan, Mr Shpiler, L. & Burra, J. 1989. The drought
R. Azzo, Mr M. Hamzeh, Mr G. Kashour, response of landraces of wheat from the
Mr A. Khorea, Mr M. Michael and Mr northern Negev Desert in Israel. Euphytica,
H. Pashayany. The work, dedication and 43: 87–96.
support of many farmers in Syria are Blum, A., Golan, G. & Mayer, J. 1991. Progress
gratefully acknowledged. achieved by breeding open-pollinated
We thank Der Bundesminister für cultivars as compared with landraces of
Wirtschaftliche Zusammenarbeit (BMZ), sorghum. Journal of Agricultural Science,
the Italian Government, the International Cambridge, 117: 307–312.
Development Research Centre (IDRC) Bramel-Cox, P.J., Barker, T., Zavala-Garcia,
and the OPEC Fund for International F. & Eastin, J.D. 1991. Selection and test-
Development for their financial support. ing environments for improved perform-
ance under reduced-input conditions. In
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275

Wolfgang Grüneberg, Robert Mwanga, Maria Andrade and Jorge Espinoza

13.1 INTRODUCTION 13.2 AN OVERVIEW OF CLONALLY

curtilobum), which evolved between important as taste, flavour and nutrient

breeders, as were their relative weighting of a breeding programme (Gabriel and

Potato (Solanum tuberosum)

Major production Quality characteristics Pest and diseases

Banana and plantain (Musa x paradisiaca)

autumnale). In the case of diploid plants breeding behaviour of allopolyploids is

after cross combinations are difficult to and doubled-triploid production. The

so that specific characteristics needed by number of genotypes at the first stage, to

are further grafted on an indicator plant certain discipline in S-clone production

Megaclones (orange and white fleshed)

Advanced breeding clones (only orange fleshed)

ELISA. The remaining 100 to 200 clones 13.8 APOMIXIS

is usually made in two or three cycles include evaluation at several locations in

by open pollination. Polycrosses have 13.12 CASSAVA

Cassava adapts to a wide range of eco- a whitefly-transmitted virus widespread in

post-harvest physiological deterioration and depression and wide segregation in cross

International Congress on Research for the International Symposium on Sweet Potato

Breeding for quantitative variables.

Daniela Soleri and David A. Cleveland

14.1 INTRODUCTION to farmers’ varieties (FVs), which include

Phenotypic selection: identification of individual plants within a population that will

Natural phenotypic selection by Artificial phenotypic selection by

Direct phenotypic selection of plants

Unintentional phenotypic selection.

Intentional phenotypic selection for Intentional phenotypic selection for

See text for key to symbols.

Copyright © 2008, D. Soleri, D.A. Cleveland, used with permission.

Conservation ( in situ in ﬁelds Improvement (via propagule

Consumption (primarily Production (primarily Multiplication (as part of

a. Traditionally-based agricultural system: functions integrated in households and

b. Industrial agricultural systems: functions separated, specialized, many institutionalized

Copyright © 2008, D. Soleri, D.A. Cleveland, used with permission.

referred to as ‘creolized’ or ‘degenerated’ the range of target environments to which

The second category emphasizes appears to have been “better understanding

definition of knowledge—both farmer and farmers continued as before. There have

phenotypically in greater trueness to farmers working in combination with

1990; Kloppenburg, 1988) and Switzerland industrial agriculture, and is expanding in

availability of human-managed inputs, such mental variation is minimal, there may be

ties may be significantly related to ethnic- of E2 and G2 to P2 , and if they distin-

Seed from long ears only is planted

Phenotypic selection for a given trait

S≈0, Phenotypic selection S>0, Phenotypic selection

R≈0, Phenotypic selection R>0, Phenotypic selection

E≈0, Maintenance or E>0, Directional or

See text for key to symbols.

or micro-evolution (E) [hereafter referred of later generations, resulting from

ly acknowledge the UCSB Faculty Senate, extent and implications. In [Proceedings

J., Martini, A.M., Salahieh, H., Goodchild, Anthropology, 38: 477–515.

Swift, M.J. 1997. Agricultural intensifica- US National Plant Germplasm System.

Breeding for quantitative variables

15.1 INTRODUCTION The switch from amateur to professional

15.1.2 Plant breeding break down as new races, strains, biotypes

ance of choice, and horizontal resistance Being quantitative, in contrast, horizontal

by a matching strain, against which the The misleading effects of parasite

resistance will appear to have decreased, cannot be observed or assessed. Because

increased epidemiological competence. 15.3.7 Biological anarchy