Professional Documents
Culture Documents
Reynolds,
J.I. Ortiz-Monasterio,
and A. McNab,
Editors
A pplication of
Physiology
in W he at Breeding
M.P. Reynolds, J.I. Ortiz-Monasterio,
and A. McNab, Editors
CIMMYT® (www.cimmyt.org) is an internationally funded, nonprofit, scientific research and training organization.
Headquartered in Mexico, CIMMYT works with agricultural research institutions worldwide to improve the productivity,
profitability, and sustainability of maize and wheat systems for poor farmers in developing countries. It is one of 16 food and
environmental organizations known as the Future Harvest Centers. Located around the world, the Future Harvest Centers
conduct research in partnership with farmers, scientists, and policymakers to help alleviate poverty and increase food security
while protecting natural resources. The centers are supported by the Consultative Group on International Agricultural Research
(CGIAR) (www.cgiar.org), whose members include nearly 60 countries, private foundations, and regional and international
organizations. Financial support for CIMMYT’s research agenda also comes from many other sources, including foundations,
development banks, and public and private agencies.
Future Harvest® builds awareness and support for food and environmental research for a world with less
poverty, a healthier human family, well-nourished children, and a better environment. It supports research,
promotes partnerships, and sponsors projects that bring the results of research to rural communities, farmers, and families in
Africa, Asia, and Latin America (www.futureharvest.org).
© International Maize and Wheat Improvement Center (CIMMYT) 2001. All rights reserved. The opinions expressed in this
publication are the sole responsibility of the authors. The designations employed in the presentation of materials in this
publication do not imply the expression of any opinion whatsoever on the part of CIMMYT or its contributory organizations
concerning the legal status of any country, territory, city, or area, or of its authorities, or concerning the delimitation of its
frontiers or boundaries. CIMMYT encourages fair use of this material. Proper citation is requested.
Printed in Mexico.
Correct citation:Reynolds, M.P., J.I. Ortiz-Monasterio, and A. McNab (eds.). 2001. Application of Physiology in Wheat
Breeding. Mexico, D.F.: CIMMYT.
AGROVOC Descriptors:Plant breeding; Breeding methods; Plant physiology; Wheats; Germplasm; Selection;
Environmental factors; Resistance to injurious factors; Yields
Additional Keywords: CIMMYT
AGRIS Category Codes: F30 Plant Genetics and Breeding
F60 Plant Physiology and Biochemistry
Dewey Decimal Classif.: 631.53
ISBN: 970-648-077-3
Design and layout: Marcelo Ortiz S., Eliot Sánchez P., and Miguel Mellado.
Cover photo credits: Demonstration of the infrared thermometer, Poza Rica, Mexico, G.P. Hettel
Waterlogging treatment, Ciudad Obregon, Mexico, K.D. Sayre
Wheat on the edge of the Sahara Desert, Egypt, M.P. Reynolds
C O N TE N TS
iv Preface
iii
PREFA CE
We applaud this practical guide to the application of physiology in wheat breeding, which
brings together in one volume the working knowledge of a broad range of experts in salinity,
drought, cold, waterlogging, micronutrients, and other key topics.
The more understanding plant breeders have of the physiological processes that underlie plant
performance, the more efficiently they can exploit relevant physiological mechanisms to
improve crop performance. Wheat breeders have become increasingly able to use physiological
traits directly as selection criteria, as their knowledge of physiological processes has expanded
and as traits have been identified that can be used as selection criteria to achieve results more
quickly and efficiently than selecting for yield performance alone.
Nonetheless, there are still major gaps in our understanding of how crops adapt to the
environment, and this calls for further physiological research. Indeed, a more complete
understanding of crop physiology will be a prerequisite to the effective application of new
techniques such as genetic transformation, functional genomics, and marker-assisted selection
in wheat breeding.
The improved varieties developed though wheat breeding are important catalysts for increasing
crop performance at the farm level, where a range of biotic and abiotic stresses impinge on
yields. However, for the maximum genetic yield potential of improved varieties to be fully
expressed, scientists must also pay due attention to crop management practices. Without
adequate soil fertility, appropriate planting methods, effective control of weeds and pests, and
efficient water management, the full economic benefits of genetic improvement can never be
realized.
Brief theoretical explanations are provided throughout this book, but the main focus is on
practical procedures breeders can readily apply. Such topics as economic issues related to the
role of physiology in wheat breeding and the search for genetic diversity that could contribute
to increasing yield will help breeders take full advantage of existing methodologies and
resources to do their work more efficiently. The chapter on the genetic basis of physiological
traits brings out the point that though field testing is indispensable, proper combination with
molecular data could lead to more efficient use of limited resources.
The collected wisdom contained in this book was generously contributed by the authors, and we
thank them for sharing the fruits of their varied experience. Through this book, their expertise
will be accessible to breeders everywhere, but especially in developing countries, where
information on this newly emerging field is rarely available.
____________________________ ____________________________
Sanjaya Rajaram Norman E. Borlaug
Director Senior Consultant
CIMMYT Wheat Program CIMMYT
iv
GENERAL CONSIDERATIONS
IN PHYSIOLOGICAL
BREEDING
1
I N TR O DU CTI O N
Application of Physiology
in W heat Breeding
M.P. Reynolds, R.M. Trethowan, M. van Ginkel, and S. Rajaram1
How can disciplinary research in their obvious phenotypic expression (i.e. A recent survey of plant breeders and
physiology complement wheat earliness versus lateness) has permitted physiologists addressed the question of
breeding? This introductory chapter is them to be modified in many breeding how physiological approaches in plant
intended to provide broad guidelines to programs. The same is true for the breeding could have greater impact
help breeding programs: 1) assess height reduction (Rht) gene. In the future (Jackson et al., 1996). According to the
whether physiological criteria should be an increased understanding of the survey, while the impacts of
included in a breeding strategy; 2) genetic basis of these traits may enable physiological research on breeding
evaluate specific physiological selection breeding programs to exploit them programs have been limited in the past,
traits and determine their usefulness in further. future impacts may arise through:
breeding. The other chapters in this
Selection for reduced height and • Focusing physiological work on an
book provide more explicit information
improved adaptation to environment has appropriate range of germplasm
on how physiological approaches can be (which will depend on the specific
had a profound impact on modern plant
used in breeding work for a variety of breeding objectives);
breeding, and the improvement in yield
environmental conditions. • Working with larger populations to
potential of spring wheat since the
enable extrapolation of findings to
Physiological criteria are commonly Green Revolution has been shown to be
breeding methods;
though not explicitly used in breeding associated with a number of other
• Identifying traits for use as indirect
programs. A good example is selection physiological factors (Reynolds et al., selection criteria, in addition to those
for reduced height, which improves 1999). Nonetheless, most breeding already used in core breeding
lodging resistance, partitioning of total programs do not put much emphasis on programs;
biomass to grain yield, and selecting physiological traits per se • Identifying traits for use as selection
responsiveness to management. Another (Rajaram and van Ginkel, 1996). criteria in introgression programs;
is differential sensitivity to photoperiod Exceptions would include: 1) the stay- • Conducting selection trials in more
and vernalizing cold, which permit green character, which has been selected representative environments, and
adaptation of varieties to a wide range for in relation to improved disease • Developing tools that could be
of latitudes, as well as to winter- and resistance and is associated with high quickly and easily applied to large
spring-sown habitats. Despite a lack of chlorophyll content and photosynthetic numbers of segregating lines.
detailed understanding of how rate in Veery wheats, for example Seri-
In this and the following chapters, many
photoperiod and vernalization 82 (Fischer et al., 1998), and 2) more
of these suggestions are incorporated
sensitivity interact with each other and erect leaf angle, a common trait in many
into a research framework for assessing
the environment, the relatively simple high yielding bread and durum wheat
the value of physiological selection traits
inheritance of photoperiod (Ppd) and plant types that was introgressed into the
in a breeding context.
vernalization (Vrn) sensitivity genes and CIMMYT germplasm pool in the early
1970s (Fischer, 1996).
1 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
2
Assessing the breeding approach will achieve results
Potential of more quickly and efficiently than
Physiological Selection selecting parents and/or progeny for
performance alone.
Criteria to Complement High spike photosynthesis
Breeding Strategies Many traits may appear to be of potential
benefit to yield. To assess which trait(s)
Stem reserves
The Art, Science, and should be prioritized, alternate
Empiricism of Breeding hypotheses may be tested empirically,
Breeding is frequently referred to as a based on a conceptual model that
blend of science and art, as well as an incorporates current understanding of Cellular traits: osmotic adjustment,
heat tolerance, ABA, etc.
empirical process. The science refers to physiological and biochemical
the routine application of established constraints to performance (Figure 1).
facts, such as the documented role of For example, if the principal yield-
specific genes in conferring disease limiting factor is water stress, Leaf traits: wax, rolling, thickness, etc.
resistance or environmental adaptation. physiological understanding would
The art of breeding refers to the intuition suggest that genotypes with deeper roots High pre-anthesis biomass
gained by working with germplasm and will have an advantage over others,
the integration of that experience with assuming moisture is available deeper in Early ground cover
established knowledge. In other words, the soil profile. However, selection for Long coleoptile
intuition enables good breeding decisions yield alone will not guarantee that good
Large seed
to be made based on our incomplete lines have the deepest roots, because
knowledge of the biology and ecology of drought tolerance may be conferred
through genetic superiority of other H2 O
plants. Empiricism refers to the use of Water relations traits:
multiple crossing and selection strategies mechanisms, such as osmotic adjustment, stomatal conductance, etc.
to achieve a single objective, sometimes accumulation and remobilization of stem
loosely referred to as “the numbers reserves, superior spike photosynthesis, Figure 1. A conceptual model for drought
game.” Physiological understanding adds heat tolerant metabolism, and good tolerance in wheat. The theoretical ideotype has
to the science, and as such complements emergence and establishment under high expression of the following traits (not all of which
the intuitive knowledge required to moisture stress (Figure 1). would be useful in all drought environments): seed size
conduct good breeding. Use of & coleoptile length (improve early crop establishment),
Genetic advance in dry environments has
physiological selection criteria can early ground cover & pre-anthesis biomass (reduce
been quite limited in most breeding evaporation of soil moisture), stem reserves /
improve the probability of success by
programs. Slower progress in moisture remobilization & spike photosynthesis (help grainfilling
making empirical selection more
stressed environments compared to during severe post-anthesis stress), stomatal
efficient.
irrigated environments is usually ascribed conductance (indicative of roots which are able to
to the heterogeneity of selection nurseries extract soil water at depth), osmotic adjustment
Theoretical Basis for Using (maintains cell functions at low water potential),
under dry conditions, which renders
Physiological Traits accumulation of abscisic acid (pre-adapts cells to
performance-based selection unreliable.
Assuming significant genetic diversity for stress), heat tolerance (heat stress may be caused by
Selection for specific traits in more
a trait is established, the question of how low leaf transpiration rates under drought), leaf
controlled environments is likely to be
its use as a selection criterion would anatomical traits e.g. waxiness, pubescence, rolling,
more effective. In addition, if more than
improve breeding efficiency can be thickness (reduce risk of photo-inhibition), high tiller
one mechanism is involved in drought
addressed. Without experimentation, this survival and stay-green (easily observed integrative
tolerance, deliberate selection with a
cannot be predicted with any certainty, traits indicative of good drought tolerance).
view to combining synergistic traits is
any more than a breeder can know in
likely to achieve results sooner than
advance which of many crosses will
adopting a strategy whereby parents and
produce the desired variety. An essential
progeny are selected on the basis of
question is whether selection for a given
performance alone (see chapter by
physiological trait as part of an integrated
Richards et al.).
0.50
R = ih2σp (1) y = 3E-09x2 + 0.0001x – 0.4153 Table 5. Criteria for applying physiological
r2 = 0.56 traits in a breeding program.
where i = intensity of selection or the 0.40
proportion of the population included in
10 12 14 16 18 20 22 24 Association of trait with
Biomass (t ha-1) Trait performance
the selected group and σp is the
phenotypic standard deviation. Figure 2. Relationship between spectral heritability Strong Weak
reflectance index (NDVI, normalized Low Selection in PYTs No application
In this case, realized heritability difference vegetation index) measured
(estimated as σ2g/σ2p) is perhaps the most during grainfilling and biomass of irrigated High Early and/or Negative selection
spring wheat advanced lines, Obregon, late generation in early
northwestern Mexico, 1996-97. selection generations
Source: Reynolds et al. (1999).
Plant breeding has traditionally applied a Given the success of such approaches to Applying Physiological
trial-and-error approach in which large date, to what extent can plant breeding Understanding in
numbers of crosses are made from many programs benefit from physiological Breeding
sources of parental germplasm. research? A recent survey of plant
Progenies are evaluated for characters of breeders and physiologists (reported in Illustrated in Figure 1 are the ways in
direct economic interest (e.g., grain yield Jackson et al., 1996) indicated a general which physiological understanding may
and grain quality) in target view among both groups that in the be applied in a breeding program; this
environments. Good performing parental future physiological research would have provides the basis for the subsequent
germplasm, crosses, and progenies are an increasing role in plant breeding discussion. The figure also shows
selected for further use or testing. In programs. However, the same survey possible ways to enhance this process,
many programs “breakthroughs” in also indicated that many respondents felt all of them based on physiological
improvement are made simply by outputs from physiological research to research or understanding.
finding superior sources of parental date had not developed into practical
germplasm among the numerous sources improvements to the extent they had Understanding yield-
tested. This conceptually simple expected or thought was desirable. limiting factors
approach has been highly successful in Understanding the biological factors
There has been considerable discussion limiting the performance of genotypes
many crop species and numerous
in the literature about the potential role across target environments is essential
breeding programs.
of physiological research in plant for improving breeding programs
The approach has often succeeded in the breeding. Much of this discussion has through physiological research.
absence of in-depth knowledge about the been from a physiological perspective— Examples of such factors are moisture
physiological basis for superior i.e., examining the potential merits of stress during different phenological
performance. In some crops such different plant traits for improving yield periods (Fischer, 1979; Woodruffe and
knowledge has been obtained by doing under different environmental Tonks, 1983), soil fertility constraints
retrospective analyses of prior genetic conditions. The aim of this chapter is to (Carver and Ownby, 1995), production
gains. Breeders have not applied this take a breeding perspective of ways in of sink capacity (grain number) and
knowledge to a significant extent as a which physiological understanding may subsequent partitioning of dry matter to
guide to further improvements, but be applied in traditional breeding grain (Gallagher et al., 1975), canopy
instead have taken any avenue of approaches. The assumption is that plant light interception during reproductive
improvement that happens to arise from breeding programs will continue to rely growth (Lawn and Byth, 1974), and
direct selection for yield and economic heavily on large scale evaluation of presence of a plant disease. The defining
performance. parental and progeny populations. feature of a limiting factor in this context
Physiological understanding could is that improving genetic response to that
enhance and refine this approach. specific factor would result in higher
yields. In all physiological research
targeting crop improvement, knowledge
of what these limiting factors are for a
particular crop species x target
1 CSIRO Tropical Agriculture, Davies Laboratory, Queensland, Australia. environmental domain combination is
11
Core breeding program noted that development of crop growth
models is expensive and, in many cases,
Selecting and
crossing of parent
may not be necessary to gain an
genotypes adequate level of knowledge.
12 P.A. JACKSON
factors causing the interactions are not managed environments for selection physiological research and literature,
adequately sampled at a particular stage trials. In Queensland, Australia, wheat successful application in breeding
of selection. If there is adequate (Cooper et al., 1995) and barley programs appears rare. Possible reasons
sampling of such factors in selection (Jackson et al., 1994) lines were found for this lack of success and ways to
trials, selection should produce good to exhibit large variation in grain enhance application in breeding
results. number and grain yield under programs are discussed below.
favorable growing conditions
A stratified random sampling of Once identified, physiological traits
(influenced by water and nitrogen
environments will usually be conducted affecting response to important limiting
availability) at around anthesis. The
across the target region if limiting factors factors may be used in two ways: 1) as
responsiveness of different lines to
are not known or poorly defined. The indirect selection criteria in progeny
good conditions at around anthesis
stratification may be based on, for populations in core breeding programs,
accounted for a large proportion of
example, geographic location, soil type, and 2) to define objectives for
G×E interactions exhibited by breeding
management regimes (e.g., irrigation introgression activities (Figure 1).
populations in a sample of production
versus rainfed), or other factors that
environments. It was therefore Genetic response for yield using another
might possibly affect the relative
suggested that wheat and barley lines trait as an indirect selection criterion may
response of genotypes. This approach
should be evaluated in one or more be predicted from the following formula
will be effective to some extent;
high input (water and N) environments (Falconer and Mackay, 1996):
however, sampling of some factors may
in the early stages of selection
be deficient with such a hit-and-miss
(Jackson et al., 1994). This would
approach, and the relative weighting to CR = i hx . rg . σgy, (1)
allow effective discrimination among
apply to results from different trials
lines for adaptation across the target
during selection will be unknown. where CR is the correlated response for
environments.
yield, from selection based on character
In this situation physiological research x; i is the standardized selection
Cooper et al. (1995) tested the value of
clearly has a role to play by helping to differential (related to the proportion of
using a small number of high input
identify the significant constraints to genetic population selected); hx is the
environments for selection trials in
higher yields and determine the level of square root of the heritability for
wheat. They showed that mean yields
genetic variation for response to those character x; rg is the genetic correlation
of wheat lines across three such
constraints in particular populations. between character y and character x; and
environments had a high genetic
Using the approaches outlined in the σgy is the genetic standard deviation for
correlation with mean yields across a
previous section, physiological research character y.
larger number (16) of random
on one or a few genotypes may help
production environments. They
identify economically important The genetic correlation is the correlation
concluded that high input managed
constraints (determined as previously between genetic effects for yield and the
environments could be used at the
described). However, comparing trait used for selection, and may be
preliminary yield evaluation stage to
responses of an adequate sample of estimated from analyses of covariance.
facilitate efficient and effective
genotypes representative of those being Many physiological studies reported in
discrimination among unselected lines.
evaluated is necessary to determine the the literature have described only
This approach should provide more
extent to which the various factors elicit phenotypic correlations (i.e., correlations
reliable selection data, at a lower cost,
genetic variation. This may sometimes based on means), which may be seriously
than using a larger number of random
be done in conjunction with selection biased (either upward or downward) by
production environments for selection
trials already established in the breeding error effects or correlated environmental
trials.
program (e.g., Jackson et al., 1994). effects. Genetic correlations are more
Improved knowledge of the factors relevant than phenotypic correlations for
Using physiological traits as examining the value of traits to be used
involved in generating variation among
indirect selection criteria
materials being tested will nearly always as selection criteria.
Identification of yield limiting factors
facilitate more focused and efficient
may suggest physiological traits that Equation 1 suggests that using a trait as
selection strategies.
breeders could use as indirect selection an indirect selection criterion will be
One way this information may be used in criteria. Although this topic has effective only under rare circumstances.
breeding programs is via the use of received considerable attention in First, the heritability (i.e., the ratio of
14 P.A. JACKSON
Negative genetic correlations may exist Using physiological and planning. The donor germplasm of
between many physiological traits and understanding to define the trait being introduced will nearly
other useful characters (e.g., Misken and objectives of introgression always be inferior from an overall
Rasmusson, 1970); this may result in low programs agronomic point of view. For this reason,
or zero genetic correlations with The aim of the main steps in a core several generations of backcrossing
economically important characters. For breeding program (Figure 1) is normally toward locally adapted material and
example, many yield components are the direct development of new cultivars, selection between each generation of
negatively correlated (Adams, 1967), so and the selection of parents and progeny crossing are required to combine
that gains from selecting for one is usually based on overall estimates of adequate expression of the trait in a
component will inevitably result in a economic performance. However, in a suitable agronomic background.
decrease in other important components. more strategic phase of genetic
improvement, the breeder may select In the future, genetic engineering
Genetic linkage or pleiotropy may cause
negative genetic correlations and will parent germplasm on the basis of a approaches will increasingly be used to
either reduce the rate of progress made in specific trait such as disease resistance or provide ‘new’ genes, efficiently
introgression (in the case of linkage) or tolerance to some abiotic stress. The aim incorporate them into adapted
reduce the value of the trait being here is to introgress—i.e., introduce—a germplasm, and control their expression.
introgressed (with pleiotropy). specific trait into locally adapted The aim here is in many ways similar to
Rasmusson (1991) found pleiotropy and breeding stocks (Simmonds, 1993). that of introgression based on more
trait compensation were major factors Successful introgression of germplasm conventional breeding approaches.
limiting progress in an extensive barley from diverse outside sources has often However, the new approaches are
breeding program applying an ideotype resulted in quantum gains in potentially more powerful in that they
approach. improvement. Examples include N.E. make a wider range of genes accessible
Borlaug’s use of dwarf wheat germplasm
Thus selection for traits that could be for improving plant traits. This research
as parental material to reduce lodging,
useful as selection criteria simply on the will require an important and
and the use of Saccharum spontaneum in
basis of physiological understanding may complementary effort by plant
sugarcane to improve ratooning and
result in small or no gains for characters physiologists and biochemists to define
stress tolerance in noble varieties
of direct interest, such as yield. Further, (Berding and Roach, 1987). or suggest specific genetic manipulations
this approach may not identify traits needed to overcome constraints to better
having a positive genetic correlation with Donor germplasm may be identified from productivity or improved quality in target
yield and in which gains via breeding any source outside the locally adapted environments.
may be easiest to achieve. If yield itself materials being selected. For example, it
(or other characters of economic interest) may include improved materials from
is used as the key selection criterion, other breeding programs which, though
physiological traits influencing yield, and not locally adapted, may have desirable Conclusions
for which genetic variation exists, will characters; materials from germplasm This chapter outlines three ways in which
automatically be changed. collections of the same species; or knowledge developed from physiological
materials from related species. research may be used to assist plant
In summary, when considering the use of Physiological understanding may be
physiological traits as indirect selection breeding programs:
useful or even necessary for choosing
criteria, expected results should be donor material to be used in introgression. • to improve sampling of environments
compared with predicted gains using Its role here would be to define specific for selection trials;
yield itself. The search for, and traits or responses of particular value for • to identify traits that may be used as
assessment of, traits that may be useful as indirect selection criteria, most
which little genetic variation exists in core
selection criteria should be based on commonly in an index combined with
breeding populations (see chapter by
estimation of their heritability, their direct measurements of economic
Skovmand et al.). performance;
measurement cost, and their genetic (not
phenotypic) correlation with yield. Using It should be emphasized that • to assist in determining the objectives
introgression is sometimes a difficult, of introgression programs and,
traits in association with yield as a
increasingly in the future, of genetic
combined selection index should also be long-term process and often unsuccessful;
engineering approaches.
considered. it therefore requires careful consideration
16 P.A. JACKSON
C HAPTER 2
Searching Genetic Resources for
Physiological Traits with Potential
for Increasing Yield
B. Skovmand,1 M.P. Reynolds,1 and I.H. Delacy2
World demand for wheat is growing at improvement. Nonetheless, many traits Genetic Resources
approximately 2% per year (Rosegrant et reportedly have potential to enhance
al., 1995), while genetic gains in yield yield, and high expression of these can The genetic resources available to plant
potential of irrigated wheat stand at less be found in germplasm collections. Seed physiologists and breeders are found in
than 1% (Sayre et al., 1997). Thus global multiplication nurseries can be used for several Triticeae genepools recognized
demand for wheat is growing at about characterizing and evaluating germplasm by Von Botmer et al. (1992) and
twice the current rate of gain in genetic collections for non-disease and non- described as concentric circles (Figure
yield potential, with progress in rainfed destructive traits (DeLacy et al., 2000). 1). The concept of the genepool was
environments being even lower. Meeting Since seed regeneration activities are first proposed in 1971 by Harlan and
these demands by continuing to expand carried out anyway, they can be an deWet (Harlan, 1992), who suggested a
agricultural production into remaining economic way of collecting data. Recent circular way of demonstrating the
natural ecosystems is environmentally work (Hede et al., 1999; DeLacy et al., relationships among genepools. The
unacceptable, and the economic costs of 2000) has indicated that agronomic traits primary genepool consists of a given
increasing yields through the (including those with low heritability) biological species including, in the case
intensification of agronomic measured on small, seed-increase of a crop species, its cultivated, wild,
infrastructure are high. Hence a cost- hillplots or miniplots can be used for and weedy forms. Gene transfer within
effective and environmentally sound such purposes. species of the primary genepool is not
means of meeting global demand for
grain is through the genetic improvement
of wheat.
17
difficult. Table 1 lists the diploid, grasses from which gene transfer can centers involved in genetic resource
tetraploid, and hexaploid species in only be achieved through the use of conservation and utilization. The
cultivated wheat’s primary genepool, special techniques. The genera and categories are:
listing their common names and species in the tertiary genepool,
indicating the genomes of each. The described by Dewey in 1984, are too
• modern cultivars in current use
• obsolete cultivars, often the elite
secondary genepool comprises the numerous to be listed here. cultivars of the past, many of which
cenospecies, i.e., a group of related are found in the pedigrees of modern
Genetic resources of cultivated plant
species with which gene transfer is cultivars
species were categorized by Frankel
possible but difficult. Most species in • landraces
(1977) and the Food and Agriculture
wheat’s secondary genepool, along with
Organization of the United Nations
• wild relatives of crop species
their synonyms and genomic • genetic and cytogenetic stocks
(FAO) Commission on Plant Genetic
constitution, are given in Table 2. • breeding lines
Resources (FAO, 1983); however, this
Wheat’s tertiary genepool is composed Recently the International Plant Genetic
categorization is not followed by all
of related genera of annual and perennial Resources Institute (IPGRI) and FAO
jointly developed a list of multi-crop
passport descriptors to provide coding
Table 1. Summary of taxa in the primary and secondary genepools of cultivated wheat. schemes consistent across crops. These
descriptors should be compatible with
I. Sect. Monococcon Dumont; Ploidy level: diploid; genome type (female x male parent): AA (‘A’) future IPGRI crop descriptors and with
1. Triticum monococcum L. those used by the FAO World
a. ssp. monococcum; cultivated form; einkorn or small spelt wheat Information and Early Warning System
b. ssp. aegilopoides (Link) Thell.; wild form; synonym: T. boeoticum (WIEWS) on Plant Genetic Resources
(PGR) (Hazekamp et al., 1997). The
2. Triticum urartu Tumanian ex Gandilyan; wild form
descriptors are: 1) unknown; 2) wild; 3)
II. Sect. Dicoccoidea Flaksb.; Ploidy level: tetraploid; genome type (female [B] x male [A] parent): BBAA (‘BA’) weedy; 4) traditional cultivar/landrace;
5) breeding/research material; 6)
3. Triticum turgidum L.; cultivated and wild forms
advanced cultivar; and 7) other. Because
a. ssp. turgidu; rivet, cone, or pollard wheat
they are more generic (so as to fit multi-
b. ssp. carthlicum (Nevski) A. Loeve & D. Loeve; Persian wheat, Persian black wheat
crop classification), these descriptors are
c. ssp. dicoccon (Schrank) Thell.; emmer wheat
also less useful in single-crop
d. ssp. durum (Desf.) Husn.; macaroni wheat, hard wheat, or durum wheat
e. ssp. paleocolchicum (Menabde) A. Loeve & D. Loeve
classification systems.
f. ssp. polonicum (L.) Thell.; Polish wheat The classification system used in
g. ssp. turanicum (Jakubz.) A. Loeve & D. Loeve; Khorassan wheat CIMMYT’s wheat collection is based on
h. ssp. dicoccoides (Koern. ex Asch. & Graebn.) Thell.; wild emmer wheat the categories outlined by Frankel and
4. Triticum timopheevii (Zhuk.) Zhuk. the FAO Commission on PGR
a. ssp. timopheevii; cultivated and wild forms (Skovmand et al., 1992). Recently,
b. ssp. armeniacum (Jakubz.) van Slageren; wild form however, a list including 21 categories
was defined in the GRIP project
III. Sect. Triticum; Ploidy level: hexaploid; genome type (female [BA] x male [D] parent): BBAADD (‘BAD’)
(Skovmand et al., 2000) to describe the
5. Triticum aestivum L. biological status of materials in
a. ssp. aestivum; bread wheat CIMMYT’s collection and other genetic
b. ssp. compactum (Host) Mackey; club, dwarf, cluster, or hedgehog wheat resources. When such specific categories
c. ssp. macha (Dekapr. & Menabde) Mackey are applied to collections, utilization
d. ssp. spelta (l.) Thell.; large spelt, spelt or dinkel wheat efficiency is enhanced, making it easier
e. ssp. sphaerococcum (Percival) Mackey; Indian dwarf wheat or shot wheat for users to know exactly what they are
6. Triticum zhukovskyi Menabde & Ericz.; Ploidy level: hexaploid; genome type (female [GA] x male [A]
working with.
parent: GGAAAA (‘GAA’)
SEARCHING GENETIC RESOURCES FOR PHYSIOLOGICAL TRAITS WITH POTENTIAL FOR INCREASING YIELD 19
n
n evaluation data. In the early 1990s, the
SEARCHING GENETIC RESOURCES FOR PHYSIOLOGICAL TRAITS WITH POTENTIAL FOR INCREASING YIELD 21
germplasm banks have traditionally agronomic, and grain quality attributes Since not all the variation is modeled,
avoided these traits, which are useful for as part of the routine regeneration some distortion of the relationships
plant improvement programs. process conducted by the CIMMYT among attributes and accessions will
Descriptions of germplasm based on Wheat Genetic Resources Program. A occur when depicted on the biplot.
“useful” attributes are immediately pattern analysis (combined use of
The attribute vectors are drawn in a
advantageous to practical plant classification and ordination methods) of
positive direction, i.e., in the direction of
improvement programs because they the data provided a good description of
increasing value for that attribute. The
indicate where useful variation may be the accessions and collection sites
length of each vector is proportional to
found in the collection. (Figure 3). Since economically useful
how well each attribute was modeled,
attributes were used, the analysis
A description based on useful attributes since each vector should be the same
provided relevant information for both
also allows more directed search
germplasm curators and potential users,
strategies than those derived from
who now have a description of the
traditional characterization attributes or Vector 2 (13.7%)
accessions from which to choose
random DNA markers with high 0.2
relevant breeding material.
heritabilities. Provided that random Attribute vectors
markers adequately cover the genome, The data were analyzed using range
they give information on the amount of standardized squared Euclidean distance 0.1
variation at and between sites, thus (rsSED) as the dissimilarity measure. Flw Mat Pr%
NoSpikes
indicating whether adequate collection These SEDs are calculated among
has been done. However, until adequate attributes that are range standardized 0.0 GrNo / S
GrWt / S
HI
linkages to known functions have been (Williams, 1976) and employed to
established, they, like traditional ensure each attribute contributes equally
-0.1 SDS
characterization attributes, provide little to the analysis. Ordination was
Hard
information on the type of variation performed by singular value
present. decomposition (Eckart and Young, 1936) -0.2
of the Gower complement similarity -0.2 -0.1 0.0 0.1 0.2
When low(er) heritability data are used,
measure to the rsSED (Gower, 1967; Vector 1 (25.4%)
means and variances change in different
DeLacy et al., 1996). The relationships
seasons, years, and places. This has
between accessions and attributes from Vector 2 (13.7%)
limited their use for germplasm 0.2
the ordination were displayed on a biplot
description, but many, if not most, Accessions and attribute vectors
(Gabriel, 1971).
attributes useful for plant improvement
programs are of this type. Much of the Both the accession and attribute plotting 0.1
difficulty encountered in integrating points can be interpreted as vectors on
such information from sets of data the biplot, but since the accessions were
acquired at different times can be investigated in terms of the attributes, 0.0
avoided by appropriate data analysis. attributes were represented as vectors
After standardization by the range or and accessions as points. As the data
-0.1
standard deviation within sets, means were centered (i.e., the attribute mean
and variances for each attribute are was subtracted from all values for that
the same. attribute so the grand mean becomes -0.2
zero), the origin on the biplot represents -0.2 -0.1 0.0 0.1 0.2
As an example, DeLacy et al. (2000)
average values for all attributes. The Vector 1 (25.4%)
reported on an analysis of a seed
percentages on each axis represent the
multiplication nursery made up of 465 Figure 3. Attribute vectors and accession
proportion of total variation, measured
accessions of bread wheat landraces plotting points for the biplot for vectors 1
by the total sum of squares (TotSS), and 2 from the ordination based on 15
collected in 1992 from 24 sites in three
accounted for by each vector. In this case morphological, agronomic, and grain quality
states of Mexico. They were examined
the aim is to represent the original 15 attributes of 465 individual spike accessions
in unreplicated hillplots in a
dimensional space defined by the 15 of wheat landraces collected from 24 sites
screenhouse for 15 morphological,
attributes in a low dimensional space. covering four states in Mexico.
SEARCHING GENETIC RESOURCES FOR PHYSIOLOGICAL TRAITS WITH POTENTIAL FOR INCREASING YIELD 23
markers could be used to screen low yield, and highly specific results; moreover, the first is
untapped genetic stocks, enabling new environmental adaptation, in addition to progressive, while the second is
sources of genes with potentially useful the needed trait. unprogressive in terms of yield potential.
alleles to be exploited in breeding.
These resources therefore need to undergo The open-parent, cyclical crossing
prebreeding before they can be used in program described by Rasmusson (2001)
How to use the
improvement work. Figure 4 demonstrates is utilized when the introgressing a trait
identified traits
two prebreeding schemes, each with a known to be of value. Rasmusson was
Genetic resources with desirable traits
different purpose: the open-parent, striving to introgress characters from
usually need to be tested and improved
cyclical crossing program and a two-row barley into six-row barley and
to be of use in wheat improvement
backcrossing program aimed at producing found that the initial cross yielded
(Figure 2). Most often these resources
isogenic lines. These two program have germplasm with no putative candidates
have many undesirable characteristics,
different purposes and different end for cultivar release, with the best lines
such as extreme disease susceptibility,
yielding about 20% less than the
improved parent. The second cycle of
the program, where the improved parent
was the best current cultivar, produced
progenies that yielded about 98% of the
best parent’s yield. The third cycle, again
Open Parent Backcrossing using the best current cultivar as a
Cyclical Crossing to recurrent parent, yielded 112-119% of the checks.
Program† parent Using this scheme, germplasm with the
desired trait is produced that could be
GR x Parent 1 (Best cultivar at the time) GR x Rec Parent
competitive in a cultivar-release
program.
Selection cycle 1 F1 x Rec Parent
A backcrossing program to generate
Selection cycle n Select plant of BC1F1 x Rec Parent isogenic lines is applied when the
identified trait has as yet no proven
Yield test selection of
value. The recurrent parent is crossed
Advanced Line 1 (AL1)
repeatedly to the genetic resource with
AL 1 x Parent 2 (Best cultivar at the time) Select plant of BC2F1 x Rec parent the desired trait. In each backcross
generation, selection is done for the tails
Selection cycle 1 Select plant of BC3F1 x Rec parent of the populations, i.e., lines with the
trait and lines without the trait. Lines
Selection cycle n Select plant of BC4F1 x Rec Parent that differ genetically only for the trait in
question are the end result of this
Yield test selection of program. Additional trials can be
Advanced Line 2 (AL) conducted to assess the value of the trait,
but bearing in mind that the germplasm
AL 2 x Parent 3 (Best cultivar at the time) Select plant of BC5F1 x Rec Parent produced will not outperform the
recurrent parent.
Selection cycle 1 Selection cycle 1
SEARCHING GENETIC RESOURCES FOR PHYSIOLOGICAL TRAITS WITH POTENTIAL FOR INCREASING YIELD 25
Table 7. Expression of the multi-ovary trait and yield components in F1 lines, wheat germplasm bank accessions, although
screenhouse, Mexico, 1999. laboratory protocols are required for
Yield component: Kernel Kernels / Kernel Grain wt / their identification. High spike
Line / cross no. / spike floret wt (mg) spike (g) photosynthesis is another trait that could
contribute to yield under stress but
Multi-ovary line 124.0 2.17 37.5 4.65 which is very time consuming to
Pastor 69.3 1.00 51.5 3.57 measure. For traits that are difficult to
Multi-ovary line/Pastor 125.9 1.81 42.1 5.30
measure (and/or that show marked
Pastor/Multi-ovary line 108.5 1.65 45.0 4.88
genotype by environment interaction), it
Baviacora M 92 72.7 1.00 57.5 4.18
Multi-ovary line/Baviacora M 92 84.6 1.04 62.8 5.31 is logical to develop genetic markers,
Baviacora M 92/Multi-ovary line 73.5 1.03 60.0 4.41 which can be used to confirm their
Esmeralda M 86 95.3 1.00 53.0 5.05 presence more unequivocally than by
Multi-ovary line/Esmeralda M 86 91.8 1.13 59.1 5.43 measuring phenotypic expression.
Esmeralda M 86/Yanglin 96.3 1.20 53.8 5.18
Conclusions
has been shown to be associated with thickness), high tiller survival, and stay- The last 30 years have witnessed an
increased leaf photosynthetic rate and green (Reynolds et al., 1999). unprecedented level of international
higher yield in improved durum wheat CIMMYT’s germplasm collection is wheat germplasm exchange and the
cultivars grown under irrigated conditions being screened, as resources allow it, for development of a greater degree of
(Pfeiffer, pers. comm.). These two high expression of many of these traits. genetic relatedness among successful
findings suggest that combining higher cultivars all over the world. The concept
High stomatal conductance permits leaf
chlorophyll content with greater spike of broad adaptation has thus been well
cooling through evapotranspiration; this,
fertility (for example, due to multi-ovary vindicated. However, greater genetic
along with higher leaf chlorophyll
florets), which creates higher demand for relatedness is seen by some as increasing
content and stay-green, is associated with
photosynthesis, may help increase yield genetic vulnerability to pathogens,
heat tolerance (Reynolds et al., 1994).
potential under irrigated conditions. although such vulnerability depends
Recent studies identified high expression
more on similarities in resistance genes,
of these traits in bank accessions, and
Traits to raise yield under which may actually be more diverse now
both traits showed high levels of
stress conditions than before. Various new factors
heritability under heat stress
Wheat yields are reduced by 50-90% of (including the growing strength of
(Villhelmsen et al., 2001). As a result,
their irrigated potential by drought on at national breeding programs in the
these accessions are currently being
least 60 million ha in the developing developing world and the advent of
crossed into good heat tolerant
world. At CIMMYT attempts are breeders’ rights) should result in
backgrounds.
underway to improve drought tolerance increased diversity among cultivars and
by introgressing stress adaptive traits into Pubescence and glaucousness protect may lead to the exploitation of hitherto
empirically selected drought tolerant plant organs from excess radiation under overlooked specific adaptation in wheat.
germplasm. Our current conceptual stressful conditions (see Loss and
This would be especially important if
model of a drought resistant cultivar Siddique, 1994). Searches are under way
climate change accelerates. Just as
encompasses high expression of the for these and a number of other leaf
increasing nitrogen supply and
following traits: seed size, coleoptile traits, such as leaf rolling, leaf thickness,
improving weed control have been
length, early ground cover, pre-anthesis and upright posture, which may well play
almost universal factors driving wheat
biomass, stem reserves/remobilization, similar roles under stress.
cultivation in the last 50 years, higher
spike photosynthesis, stomatal
Osmotic adjustment (Blum et al., 1999) atmospheric concentrations of CO2 and
conductance, osmotic adjustment,
and stored stem fructans (Blum, 1998) global warming with resulting warmer
accumulation of abscisic acid, heat
have been implicated in stress tolerance. temperatures could significantly
tolerance, leaf anatomical traits (such as
Searches are underway for high influence breeding objectives in the next
glaucousness, pubescence, rolling,
expression of these traits among 50 years.
SEARCHING GENETIC RESOURCES FOR PHYSIOLOGICAL TRAITS WITH POTENTIAL FOR INCREASING YIELD 27
Loss, S.P., and K.H.M Siddique. 1994. Richards, R.A. 1996. Increasing the yield Skovmand, B., G. Varughese, and G.P. Hettel.
Morphological and physiological traits potential of wheat: Manipulating sources 1992. Wheat Genetic Resources at
associated with wheat yield increases in and sinks. In: Increasing Yield Potential in CIMMYT: Their Preservation,
Mediterranean environments. Adv. Agron. Wheat: Breaking the Barriers. M.P. Documentation, Enrichment, and
52:229-276. Reynolds, S. Rajaram, and A. McNab (eds.). Distribution. Mexico, D.F.: CIMMYT.
Maes, B., Trethowan, R.M., Reynolds, M.P., van Mexico, D.F.:CIMMYT. pp. 134-149. Tanksley, S.D., and J.C. Nelson. 1996. Advanced
Ginkel, M., and Skovmand, B. 2001. The Rosegrant, M.W., M. Agcaoili-Sombilla, and N.D. back-cross QTL analysis: A method for the
influence of glume pubescence on spikelet Perez. 1995. Global Food Projections to simultaneous discovery and transfer of
temperature of wheat under freezing 2020: Implications for Investment. valuable QTLs from unadapted germplasm
conditions. Aust. J. Plant Physiol. Washington, D.C.: IFPRI. into elite breeding lines. Theor. Appl. Genet.
28:141-148. Sayre, K.D., S. Rajaram, and R.A. Fischer. 1997. 92:191-203.
Mujeeb-Kazi, A. 1995. Interspecific crosses: Yield potential progress in short bread Trethowan, R.M., M.P. Reynolds, B. Skovmand,
hybrid production and utilization. In: A. wheats in northwest Mexico. Crop Sci. and M. van Ginkel. 1998. The effect of
Mujeeb-Kazi, and G.P. Hettel, eds. 1995. 37:36-42. glume pubescence on floret temperature in
Utilizing wild grass biodiversity in wheat Shands, H.L. 1991. Complementarity of in-situ wheat. Agronomy Abstracts p. 161.
improvement: 15 years of wide cross and ex-situ germplasm conservation from van Slageren, M.W. 1994. Wild wheats: a
research at CIMMYT. CIMMYT Research the standpoint of the future user. Israel monograph of Aegilops L. and
Report No. 2. Mexico, D.F.: CIMMYT. Journal of Botany 40:521-528. Amblyopyrum (Jaub. & Spach) Eig
Pardey, P.G., B. Skovmand, S. Taba, M.E. Van Siebeck, W.E. 1994. Intellectual Property Rights (Poaceae). Wageningen Agric. Univ.
Dusen, and B.D. Wright. 1998. The cost of and CGIAR Research: Predicament or Papers 94-97.
conserving maize and wheat genetic Challenge. CGIAR Annual Report 1993- Vilhelmsen, A.L., Reynolds, M.P., Skovmand, B.,
resources ex situ. In: M. Smale (ed.). 1994. pp. 17-20. Mohan, D., Ruwali, K.N., Nagarajan, S., and
Farmers, gene banks, and crop breeding: Skovmand, B., P.N. Fox, and J.W. White. 1998. O. Stoelen. 2001. Genetic diversity and
Economic analysis of diversity in rice, Integrating research on genetic resources heritability of heat tolerance traits in wheat.
wheat, and maize. Dordecht, The with the international wheat information Wheat Special Report No. 50. Mexico, D.F.:
Netherlands: Kluwer Academic Press. system. In: Wheat Prospects for global CIMMYT. (In preparation.)
pp. 35-55. improvement. H.-J. Braun, F. Altay, W.E. Villareal, R.L., G. Fuentes Davila, and A. Mujeeb
Pardey, P.G., B. Koo, B.D. Wright, M.E. Van Kronstad, S.P.S. Beniwal, and A. McNab Kazi. 1995. Synthetic hexaploid x Triticum
Dusen, B. Skovmand, and S. Taba. 2000. (eds.). Dordecht, The Netherlands: Kluwer aestivum advanced derivatives resistant to
Costing the Conservation of Genetic Academic Publishers. pp. 387-391. Karnal bunt (Tilletia indica Mitra). Cer. Res.
Resources: CIMMYT’s Ex Situ Maize and Skovmand, B., M.C. Mackay, H. Sanchez, H. van Commun. 23:127-132.
Wheat Collection. Crop Sci. (in press). Niekerk, Zonghu He, M. Flores, R. Herrera, Von Botmer, R., O. Seberg, and N. Jacobsen.
Rasmusson, D.C. 2001. Learning about barley A. Clavel, C.G. Lopez, J.C. Alarcon, G. 1992. Genetic resources in the Triticeae.
breeding. In: Breeding Barley in the New Grimes, and P.N. Fox. 2000a. GRIP II: Hereditas 116:141-150.
Millenium. H.E. Vivar and A. McNab (eds.). Genetic resources package for Triticum and Williams, W.T. 1976. Pattern analysis in
Mexico, D.F.: CIMMYT. pp. 1-6. related species. In: Tools for the New agricultural science. Amsterdam, The
Reynolds, M.P., M. Balota, M.I.B. Delgado, I. Millenium. B. Skovmand, M.C. Mackay, C. Netherlands: Elsevier Scientific Publishing
Amani, and R.A. Fischer. 1994. Lopez, and A. McNab (eds.). On compact Company.
Physiological and morphological traits disk. Mexico, D.F.: CIMMYT.
associated with spring wheat yield under Skovmand, B., Lopez, C., Sanchez, H., Herrera,
hot, irrigated conditions. Aust. J. Plant R., Vicarte, V., Fox. P.N., Trethowan, R.,
Physiol. 21:717-30. Gomez, M.L., Magana, R.I., Gonzalez, S.,
Reynolds, M.P., B. Skovmand, R. Trethowan, and van Ginkel, M., Pfeiffer, W., and Mackay,
W.H. Pfeiffer. 1999. Evaluating a M.C. 2000b. The International Wheat
Conceptual Model for Drought Tolerance. Information System (IWIS). Version 3. In:
In: Using Molecular Markers to Improve Tools for the New Millenium. B. Skovmand,
Drought Tolerance. J.M. Ribaut (ed.). M.C. Mackay, C. Lopez, and A. McNab
Mexico, D.F.: CIMMYT. (eds.). On compact disk. Mexico, D.F.:
CIMMYT.
During the past two decades, molecular The Genome important role in gene expression. While
tools have aided tremendously in the DNA encodes genetic information in the
identification, mapping, and isolation of Although the expression of genes can be form of messenger RNA (mRNA),
genes in a wide range of crop species. modified by environmental factors, the proteins are involved in the packaging of
The vast knowledge generated through nuclear genome of plant cells carries a DNA and in regulating its availability for
the application of molecular markers has genetic blueprint in the form of transcription. Transcribed gene products
enabled scientists to analyze the plant deoxyribonucleic acid (DNA) that are transported across the nuclear
genome and have better insight as to contains information for cell maintenance envelope to be translated into proteins
how genes and pathways controlling and replication. The nuclear genome using the cellular apparatus.
important biochemical and contains the largest amount of DNA and
the highest number of genes encoded, but Genes are distributed along the
physiological parameters are regulated.
plant cells also contain DNA in their chromosomes, and the number of
Three areas of biotechnology have had
chloroplasts and mitochondria. Nuclear chromosomes a plant cell contains varies
significant impact: the application of
genomes of crop species are estimated to among crop species. There is
molecular markers, tissue culture, and
contain thousands of genes, some unique considerable diversity in genome
incorporation of genes via plant
and others in multiple copies. However, composition and organization of different
transformation.
the amount of DNA in the nuclear genome organisms (Table 1). With the aid of
Molecular markers have enabled the represented by transcribed genes is only a molecular techniques, it has been possible
identification of genes or genomic fraction of total DNA found in the to study and understand the organization
regions associated with the expression genome. of the nuclear genome of several plant
of qualitative and quantitative traits and species. Plant genome analysis
made manipulating genomic regions Nuclear DNA is packaged and organized encompasses genome mapping, gene
feasible through marker assisted into chromosomes along with histones and tagging, quantitative trait (QTL) analysis,
selection. Molecular marker applications non-histone proteins. The interactions and synteny mapping.
have also helped us understand the between DNA and proteins play an
physiological parameters controlling
plant responses to biotic and abiotic Table 1. DNA content per haploid genome in different organisms.
stress or, more generally, those involved
Mega base pairs
in plant development. This chapter
Organism 2n Picograms † 106bp / 1C Length (cm)
discusses different types of molecular
markers, the basic principles and E. coli (1) 0.0047 4.2 0.14
practical considerations involved in their Chloroplast (maize) (c) 0.0002 0.160 0.006
application in plant improvement, and Mitochondrion (maize) (m) 0.0007 0.570 0.02
Arabidopsis thaliana 10 0.15 150 4.4
some contributions they have made to
Oryza sativa 24 0.45 430 13.1
wheat molecular genetics.
Triticum aestivum 42 5.96 5,700 173
Zea mays 20 2.6 2,500 75
Homo sapiens 46 3.2 3,900 102
† 1 picogram = 1 pg = 0.965 x 109 bp = 29 cm.
1 CIMMYT Applied Biotechnology Laboratory, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
2 John Innes Centre, UK.
29
The DNA Molecule instructions for creating a particular ensures that the zygote (after the male
In higher organisms, a DNA molecule organism with its own unique traits. The and female gametes unite) will contain
consists of a sequence of nuclear acids size of a genome is usually stated as the the same number of somatic
linked by chemical bonds. Each total number of base pairs in the haploid chromosomes as the parents.
nucleotide contains a heterocyclic ring genome (Table 1).
The chromosomal form of the nuclear
composed of carbon and nitrogen atoms
genome varies significantly during cell
(the nitrogenous base), a five-carbon Genes and Chromosomes
division. During interphase, the
sugar in ring form (a pentose), and a The gene is the basic physical and
chromatin in the chromosomes remains
phosphate group. There are two kinds of functional unit of heredity. Each gene is a
diffuse and therefore less visible under
nitrogenous bases: purines and nucleic acid sequence that carries
the microscope, it becomes more
pyrimidines. Each nucleic acid is information encoded to represent a
condensed and highly visible for
composed of only four types of bases: particular polypeptide. Polypeptides
cytogenetic manipulation during meiosis
two kinds of purines, known as adenine provide the structural components of cells
and mitosis. Cytological studies of
(A) and guanine (G), and two kinds of and tissues as well as enzymes for
individual chromosomes during
pyrimidines, cytosine (C) and thymine (T). essential biochemical functions. The plant
metaphase through chromosome
genome is estimated to comprise 20,000
The nitrogenous base is linked to the banding techniques have helped
to 100,000 genes.
pentose sugar by glycosidic bonds. characterize and identify the individual
When the phosphate group is added to Genes vary widely in length, often chromosomes of the wheat karyotype.
the pentose sugar, the base-sugar- extending over thousands of bases, but Moreover, classical cytological studies
phosphate complex is called a only about 10% of the genome is known and current molecular cytogenetic
nucleotide. Nucleotides are linked to include protein-coding sequences techniques have aided in identifying
together into a chain by a backbone (exons) of genes. Interspersed within chromosomal abnormalities and subtle
consisting of an alternating series of genes are intron sequences, which have interchanges.
sugar and phosphate residues with the no coding function. The rest of the
bases attached to the sugar molecules. In genome is thought to consist of other The Wheat Genome
higher organisms, DNA consists of two noncoding regions (such as control The numerous species of the genus
strands of nucleic acids that are wrapped sequences and intergenic regions), whose Triticum can be classified into three
around each other in antiparallel form in functions are still obscure. The ploidy groups: diploids (2n=2X=14),
a double helix. The sides of the two configuration and methyation level of a tetraploids (2n=4X=28), and hexaploids
strands are composed of sugar and DNA molecule play a role in gene (2n=6X=42). Of the Triticum species,
phosphate molecules, and the bases are expression, since expressed regions are cultivated T. aestivum, known as bread
inside the double helices. The two generally characterized by a high level of wheat, is the principal commercial type,
strands are held together by hydrogen methylation. Some genes have few whereas T. turgidum (durum wheat) is
bonding between the purine of one copies; others may be present in multiple principally used for making pasta.
strand with a pyrimidine of the opposite copies per haploid genome. Such Cultivated bread wheat is an
strand. Base A always pairs with a T via repeated sequences may be present in allohexaploid (2n=6X=42), composed
two hydrogen bonds, whereas a G tandem copies at a chromosomal locus or of three distinct genomes, A, B and D.
always pairs with a C via three hydrogen in different chromosomes dispersed Current evidence suggests that it
bonds. The composition of bases along throughout the genome. originated from natural hybrids of three
one strand of the DNA chain is exactly diploid wild progenitors native to the
complementary to its partner strand, The vast amount of DNA present in each
Middle East. Triticum urartu Tum. is
which allows both strands to carry the plant cell is tightly packaged, with the
recognized as the donor of the A
same genetic information. This is help of histone and non-histone proteins
genome. Although Aegilops speltoides
essential for the self replicating in the nucleus, into microscopic
was considered the donor of the B
capability of DNA. structures known as chromosomes. Genes
genome, current evidence suggest that
are scattered along the chromosomes,
the real donor is either extinct or an
The particular order of the bases which vary in number from species to
undiscovered species belonging to the
arranged along the sugar-phosphate species. During gametic formation the
Sitopsis section of Aegilops (Pathak,
backbone is called the DNA sequence. somatic chromosome number is divided
1940; Kimber and Athwal, 1972; Miller
This sequence provides precise genetic in half by cell division (meiosis), which
et al., 1982). Triticum tauschii, also
DNA, individual 1
primer +3
Conserved Conserved Selective
Amplified SSRs amplification with
primed region 1 primed region 2
primers +3
DNA, individual 2
Sequencing gel
Amplified SSR DNA fragments have a 6 bp difference in length, a DNA
polymorphism that can be resolved by high resolution electrophoresis EcoRI adapter sequence
Msel adapter sequence
Figure 2. Example of a microsatellite: a dinucleotide repeat showing a polymorphism
between two different individuals. Figure 3. AFLP method.
The BSA method involves screening Parent A Parent B F1 Resistant bulk Susceptible bulk
two pooled DNA samples from
individuals with contrasting traits from a
segregating population originating from BSA
a single cross. Each pool, or bulk,
contains individuals selected to have
identical putative genotypes for a
particular genomic region (target locus
or region) but also random genotypes at
loci unlinked to the selected region.
Therefore, the two bulked DNA samples Resistant Susceptible
differ genetically only in the selected
region and present random allelic
segregation for all other loci. For
RILs
example, if markers are to be identified
for disease resistance, equal amounts of
DNA from the 5-10 most resistant
individuals are bulked and taken as a Figure 8. Bulk segregant analysis.
If ≠ QTL!
Table 4. Genes identified and mapped with molecular markers for physiological and agronomic traits in wheat.
Traits Genes Species Markers Chromosomes References
Physiological and agronomic
Preharvest sprouting QTL Triticum aestivum RFLP Anderson et al., 1993
Vernalization Vrn1 RFLP 5AS Galiba et al., 1995; Korzun et al., 1997
Kato et al., 1998
Vrn3 RFLP 5DS Nelson et al., 1995a
Photoperiod response Ppd1 T. aestivum RFLP 2DS Worland et al., 1997
Ppd2 T. aestivum RFLP 2BS Worland et al., 1997
Dwarfing Rht8 SSR 2DS Korzun et al.,1998
Rht12 SSR 5AL Korzun et al., 1997
Cadmium uptake RAPD Penner et al., 1995
Aluminum tolerance Alt2 RFLP RFLP 4D 4DL Luo and Dvorak, 1996; Riede and
Anderson, 1996
Drought induced ABA RFLP 5A Quarrie et al., 1994
Na+/K+ discrimination Kna1 T. aestivum RFLP RFLP 4D 4DL Allen et al., 1995; Dubcovsky et al., 1996
Quality
Kernel hardness Ha Hn and QTL RFLP RFLP 5D 5DS, 2A, 2D, 5B, 6D Nelson et al., 1995a; Sourdille et al., 1996
Grain protein QTL T. turgidum RFLP 4BS, 5AL, 6AS, 6BS, 7BS Blanco et al., 1996
LMW glutenins T. turgidum 1B D’ovidio and Porceddu, 1996
HMW glutenins Glu -D1 -1 T. aestivum ASA 1DL D’ovidio and Anderson, 1994
Flour color RFLP/AFLP 7A Parker et al., 1998
The goal of most breeding trials is to Choosing the • Climate is an important consideration
assess the performance of a number of Experimental Site especially when looking for abiotic or
genetically diverse breeding materials in biotic stress tolerance. Should the
such a way that superior lines can be Selecting the experimental site is critical breeder select a high or low rainfall
selected that perform better than local for success. If a breeder wishes to select area, cool or hot temperatures at the
checks under specific farming conditions. materials for salinity tolerance, he must beginning or end of the growing
select a site that represents the salinity season? Is frost an important
This must be done without bias under
situation in the areas where the variety consideration or heat during
conditions that mimic the conditions
will be released. His chances of success grainfilling?
where the material will be recommended.
are increased by carefully selecting the • Is the crop being recommended for an
If the breeder is going to select material irrigated or rainfed situation? Is
for a specific biotic or abiotic stress, he site for the nursery. This selection will
waterlogging or poor drainage a
must conduct the trial in an area that has take into consideration the soil, climate, characteristic of the recommendation
this stress or condition. This will ensure a water regime, and biotic and abiotic domain? If selecting for rainfed areas,
higher rate of success than planting only stresses that will be encountered in the how do you handle the problem of
under favorable conditions that do not target environment. Factors that need to soil moisture at planting? Do you pre-
represent the target environment. be considered include: irrigate the plot so germination is
good and then leave the rest of the
It is also important to manage breeding • Soil factors such as texture, pH, season to natural rainfall? Is irrigation
trials as carefully as possible to minimize conductivity (salinity and alkalinity), water available and is it of the quality
and nutrient status. Soil texture will you need or similar to that of the
experimental error to be able to evaluate
affect soil physical properties and the target environment?
differences between materials
permeability and drainage of the soil.
statistically. Confounding factors need to • If the breeder is selecting material for
Soil pH and conductivity can have a specific biotic stresses, he should
be kept to a minimum so that the breeder large effect on plant growth, and
has confidence that his selections will select an area where this stress
crops and different cultivars will occurs. Incidence of certain diseases
stand up to testing under the client perform differently under different and insect pests comes to mind.
situation. This chapter will look at some values for these parameters. Soil
• Abiotic stresses such as heat, salinity,
of the factors that need to be considered nutrient status will affect the yield and waterlogging also need to be
when planning breeding trials. potential of the germplasm. It is also addressed when selecting a site. Are
important to consider soil nutrient abiotic stresses (for example, salinity
status when breeders are specifically or waterlogging) consistent across the
looking for tolerance to various selected field? If not, can they be
nutrient factors such as phosphorus statistically handled by experimental
efficient lines or lines tolerant to design and layout?
micronutrient deficiencies.
1 CIMMYT Natural Resources Group, P.O. Box 5186, Lazimpat, Kathmandu, Nepal.
2 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
48
Another question is whether the wheat usually follows cotton, maize, or • The previous crop will have a specific
experimental area should be on station soybeans. It is therefore important that spectrum of pests and diseases. If
or in farmers’ fields. Most breeding some breeding trials be grown on land weeds from the previous crop carry
trials have traditionally been conducted with similar cropping patterns, for over to the wheat crop, measures
on station, mainly because it gives the several reasons: must be taken to control the problem.
Residues from the previous crop can
breeder control over experimental
conditions, e.g., land preparation,
• The previous crop can strongly also influence disease and insect
influence the harvest date and incidence. In many countries
fertilization, irrigation, crop protection, therefore the planting date for the combine-harvesting is becoming
and security of the trial. However, there wheat crop. For example, in Asia, popular. This system leaves loose
is often the problem how representative long-maturing “basmati” rice is residue on the soil surface. In some
the station is of the client farmer preferred in some locations because cases, this residue can harbor
situation. Unfortunately, many stations of its high quality and market price, diseases. In others, there are benefits
are not chosen for their similarity with its straw value and the fact that it to this mulch (especially rainfed
farmers’ fields but rather for needs less fertilizer. The next wheat areas) in the form of cooler
convenience or availability. Experiment crop will inevitably be planted late. temperatures and better water
stations usually represent one or at most Since there is genetic variability for infiltration and conservation. Thus it
a few of the soils or environments faced performance of wheat varieties under is important to screen materials under
late planting, breeding trials should at these conditions.
by farmers.
some point in the selection process be
There is therefore a tradeoff between planted late after rice.
experimental control and how • The previous crop can have an effect Preparing the Nurser y
representative on-station conditions are on soil physical properties. When rice
is grown in South Asia, the soil is Unless germplasm is being selected for
of the target environment. Probably the
puddled (wet cultivation) to lower reduced or zero-tillage situations, land
best solution is to conduct breeding
permeability and water use. This preparation is a major first step in
trials under both situations. The first
profoundly affects the soil structure nursery management. The objective is to
phase selects promising materials under for the next upland crop, such as prepare the soil to favor wheat
the control of the station; later the wheat. The poor structure following germination. Several factors need to be
materials are assessed under actual rice affects rooting and soil considered:
farmer situations. The latter should permeability. Waterlogging is
preferably include farmer participation common in wheat following rice, • Selection of the appropriate plow or
and experimentation. The extra benefit with plants turning yellow due to harrow for the specific soil to ensure
of farmer participation is the feedback oxygen stress. Thus a breeder should a good tilth, favorable for wheat
and assessment he can give the breeder. evaluate his germplasm under these germination. This may involve a
The new emphasis on farmer conditions to improve the chances of series of steps from deep plowing, to
selecting adaptable varieties for rice- harrowing, to compaction of the
participatory breeding by some donor
wheat farmers. surface soil to ensure good seed-soil
and development agencies utilizes this
• Soil nutrient status is also affected. contact.
important feedback mechanism.
Some crops like rice are very • Fields should be leveled as much as
exhaustive of nitrogen. For others, possible to reduce variability due to
like potato, soil nutrient status is often soil moisture, especially if applying
Influence of Crop better than normal due to high irrigation. If germplasm is being
fertilizer application rate, which selected under rainfed conditions and
Rotations the fields are sloping, appropriate
leaves residual fertility.
In many developing countries, • Soil water status can also be affected. experimental blocking designs will be
especially in subtropical environments, Deep-rooted crops such as sugarcane needed to reduce experimental
wheat is often grown in double or triple and cotton can deplete the soil profile variability and error.
cropping patterns. For example, in Asia,
of water. If the breeder grows his • If the nursery is sown in farmers’
nursery after fallow, he will get fields, the plots should be prepared in
wheat is grown sequentially with rice,
different results than if he plants after the same way that farmers do, using
cotton, soybeans, and maize in the same the most representative areas (away
the previous crop, unless he
calendar year. In Mexico, where the compensates for the low water status from field edges, buildings, trees,
CIMMYT wheat varieties originate, by irrigation. etc.).
Table 2. Formulas for calculating yield components using three different harvesting methods.†
Harvesting method
† A=plot area harvested (m-2); SS=sub-sample; fw=fresh weight; dw=dry weight; P=plot; GW=grain weight.
Note: Formulas assume that grain dried at 70oC is at 0% moisture. Grain yield at x% moisture = Yield * [100/(100 – x)] (g/m2).
4 Using this calculation, KNO is statistically linked to GY and may give rise to spurious correlations between GY and GNO if GY is not determined
accurately.
5 This step-by-step description of the line-transect method was taken from a fact sheet by David P. Shelton, and Elbert C. Dickey, Extension Agricultural
Engineers, University of Nebraska; Robert Kanable, Conservation Agronomist, Soil Conservation Service; Stewart W. Melvin, Extension Agricultural
Engineer, Iowa State University; and Charles A. Burr, Extension Agricultural Specialist, University of Missouri. The fact sheet was produced through
the cooperative efforts of representatives of Cooperative Extension, Midwest Plant Service, NACD’s Conservation Technology Information Center,
U.S. Environmental Protection Agency, and the USDA Soil Conservation Service.
This review presents practical guidelines 1993; Loss and Siddique, 1994). leaves. However, under field conditions
for breeders of wheat (and other small However, in breeding for crop fluorescence values may only reflect
grain cereals) interested in adopting a resistance, the evaluated traits and differences in phenology across
physiological approach to crop screening tools are often related to genotypes. Nevertheless, remote sensing
improvement. Some of the most tolerance, not avoidance (see definitions detection of fluorescence spectra at the
promising tools for fast, reliable in Larcher, 1995). canopy level could become a promising
characterization of yield-determining approach for breeding purposes
An indirect (i.e., physiology-based)
traits will be discussed from an (Lichtenthaler, 1996).
breeding strategy could fail to produce
ecophysiological perspective. We will
yield gains and might even lead to a
focus on the practical aspects and
decrease in yield. Improved plant
limitations of using relevant screening
tolerance, though it protects the crop, Identifying
tools or selection criteria. The potential
contributions of physiological research
can limit yield potential. The target Physiological Traits for
to plant breeding, as well as its inherent
environment where selection is carried Use As Selection
limitations, have been extensively
out must be defined a priori, and the Criteria
possibility of a negative breeding effort
reviewed from a breeding perspective One approach to search for traits that
should not be ignored. In fact, plants that
(for example, Jackson et al., 1996). A could be used in breeding programs is to
show the most tolerant response
theoretical framework for identifying identify the physiological processes
during screening may also be the most
yield determinants that are obvious determining productivity. A crop’s yield
sensitive, in terms of yield loss, for
candidates for evaluation has also been potential (or harvestable part, Y) over a
example, because they are unable to
established (see below), although it has given period of time can be divided into
delay the effect of stress at the cellular
not been used in practical breeding. three major processes (Hay and Walker,
level.
The use of physiological traits as 1989). First, the interception of incident
The most promising methods allow for solar radiation by the canopy; second,
screening tools in breeding is still largely
quick screening of “integrative” the conversion of intercepted radiant
experimental—for different reasons.
physiological traits (Araus, 1996), so energy to potential chemical energy (i.e.,
Sometimes the traits are very indirectly
called because they integrate plant dry matter); and third, the
related to yield (Araus, 1996; Richards,
physiological processes either in time partitioning of dry matter between the
1996) or there is little ecophysiological
(i.e., during the plant cycle) or at the harvested parts and the rest of the plant.
understanding of the crop, especially
organization level (e.g., whole plant, Whereas the first component depends on
when breeding for yield under stress.
canopy). Other quick screening methods the canopy’s total photosynthetic area,
Nevertheless, breeding for crop escape
for evaluating, for example, the the second relies on the crop’s overall
has been very successful, and
photosynthetic performance of plants photosynthetic efficiency (i.e., total dry
phenological changes have been the
under stress conditions have been matter produced per unit of intercepted
most important indirect factor in
proposed—among them, chlorophyll radiant energy); the third is harvest
increasing wheat yields under
fluorescence measurements on single index. Total biomass, which is the result
Mediterranean conditions (Slafer et al.,
59
of the first two components, can be The Δ not only evaluates genotypic and external environmental conditions
physiologically defined as the result of differences in water use efficiency, but that influence photosynthetic gas
canopy photosynthesis over time. can also be affected by the total amount exchange (Farquhar et al., 1989). In C3
of water transpired by the crop (the first cereals such as wheat, Δ is positively
Other approaches may be followed, related to CO2 levels in intercellular air
component of Passioura’s equation) or
depending on agroecological conditions. spaces (Diagram 1) (Farquhar et al.,
by photosynthetic activity (the second
For example, under water-limiting (e.g., 1982; Farquhar and Richards, 1984;
component of the yield potential
Mediterranean) conditions, the most Ehdaie et al., 1991) and, given a constant
equation). Keeping in mind developing
widely used framework, proposed by leaf-to-air vapor pressure difference, also
country breeders, we have included
Passioura (1977), allows the study of negatively related to water use efficiency
under the generic title “surrogates” other
indices that maximize yield per rainfall (WUE, measured either as net
screening criteria, such as ash
unit. Thus economic yield depends on photosynthesis/transpiration, also called
accumulation in different plant parts or
the total water transpired by the crop, transpiration efficiency, or as plant
criteria related to leaf structure. Though
water use efficiency, and harvest index. biomass produced/water transpired)
these features are not Δ surrogates in a
Although these three components are not (Farquhar and Richards, 1984; Hubick
strict sense, they are always related to
truly independent, Passioura’s is a useful and Farquhar, 1989). Plants with high
yield and are both quicker and cheaper WUE would be less able to discriminate
framework for searching for critical
than Δ determinations; furthermore, no against 13C, and thus would accumulate
traits to improve yield under drought.
large facilities or highly qualified more of the heavy carbon isotope in their
Yield can be divided into several technical support are necessary to tissues than less efficient water users.
integrative components or traits. Yield use them.
itself is the most integrative trait, When measured in plant dry matter, D
Canopy spectral reflectance is one of the provides an (integrated) indication of
because it is influenced by all factors
most promising remote sensing WUE throughout plant growth (Farquhar
(known and unknown) that determine
techniques (see also Araus, 1996). et al., 1982, 1989). D has been proposed
productivity. However, there are many
Although at present the equipment is as a possible screening tool for
known limitations in a purely empirical
very expensive, in a few years its cost identifying variations in WUE in wheat
breeding approach based only on yield.
should drop dramatically. Another (Farquhar and Richards, 1984; Ehdaie et
Therefore, any breeding strategy based
remote sensing technique, canopy al., 1991; Condon and Richards, 1993)
on a physiological (i.e., analytical) and barley (Hubick and Farquhar, 1989).
temperature, provides integrated
approach should use screening tools or In fact, the permanent relationships
information on the crop’s stomatal
criteria to evaluate the integrative between WUE and D during treatment
conductance at the canopy level (see
physiological parameters that determine and the high broad-sense heritability of
chapter by Reynolds) (Reynolds et al.,
harvestable yield with a single D in wheat, together with other
1994) and has the advantage of being
measurement. Although harvest index considerations, indicate that D may be
low cost. However, its usefulness is
has been the most successful trait when useful for modifying the WUE and yield
limited in severely stressed
modified to improve yield, the other two of water-limited wheat crops (Condon et
environments, and genotypic differences
components of the above equations, al., 1987; Condon and Richards,
in phenology and canopy architecture
which are responsible for total crop 1992, 1993).
can further limit its validity.
biomass, remain (basically) unchanged.
In the following pages we will focus on The relationship between Δ
tools used to evaluate physiological traits and water-use efficiency
determining total biomass. We will Carbon Isotope Following on the model of Farquhar et
discuss two different kinds of tools for Discrimination al. (1982), Δ in C3 plants may be defined
screening integrative physiological traits: in its simplest form as:
For C3 plants, discrimination (Δ) of the
carbon isotope discrimination (Δ) and
heavier (13C) stable carbon isotope over
indices based on canopy spectral- Δ = a + (b - a)(pi /pa),
the lighter, more abundant (12C) form
reflectance.
(99%) during photosynthetic CO2
where a is the carbon-13 discrimination
fixation is an integrated measure of
caused by diffusion in air (4.4 0/00), b is
internal plant physiological properties
that caused by carboxylation by the
Δ = (δa - δp Δ / (1 + δp )
20 18.5
r2 = 0.17 y = -3.67 + 0.62x a r2 = 0.86 a
r2 = 0.21 y = -10.12 + 1.19x
r2 = 0.27 y = -10.42 + 1.44x 17
Δ Kernels ( 0/ 00 )
15
Ash flag leaf (% dry mass)
15.5
13 14 15 16 17 18
10
c
2.2
1.6
Ash kernels (%)
1.8
1.4
1.2
r2 = 0.28 y = 1.44 - 0.000167x
1.0 r2 = 0.19 y = 1.91 - 0.000111x
r2 = 0.22 y = 2.03 - 0.000057x
0.8 0.6
13 14 15 16 17 18 1500 3000 4500 6000 7500 9000
Figure 2. Relationship between carbon isotope discrimination (Δ) in Figure 3. Relationship between grain yield and (a) carbon
mature kernels and ash content based on dry mass (a) of the flag leaf isotope discrimination (Δ) in mature kernels, (b) ash content
blades around three weeks after anthesis, and (b) in the same mature (based on dry of the flag leaf around three weeks after
kernels. Plants were cultivated in three trials differing in water status: anthesis, and (c) ash content of the same mature kernels.
Breda, Tel Hadya rainfed, and Tel Hadya with supplementary irrigation.
Figure 4. Reflectance signature of two wheat leaves differing in nitrogen status. Note the
higher reflectance in the photosynthetically active radiation region of the nitrogen deficient
leaf due to lower chlorophyll content in the leaf area.
Measurement techniques
Instruments. The instruments required
for measuring reflectance spectra are: 1)
a field spectroradiometer that analyzes
the spectrum of sampled radiation, 2)
foreoptics that capture the radiation
reflected by a given target, and 3)
reference panels, supports, and levels for
repeated sampling of incident radiation
and radiation reflected by the canopy.
Wheat breeding can be thought of as an are established, the informal and ad hoc to increase protein content in wheat,
economic activity, in the sense that it manner in which this is done frequently experience suggests that it will often be
involves processes of physical leads to decisions that are far from better to concentrate on breeding for
transformation that are characterized by optimal in an economic sense. Basic higher yield, while leaving the challenge
streams of costs and benefits. Decisions economic analysis, grounded in the of raising protein content to agronomic
taken about the organization and careful assessment of benefits and costs, management. This is because even
operation of a wheat breeding program can provide the foundation for making though cultivars differ in their protein
(including technical decisions such as those decisions in a more informed and content, prospects for increasing protein
the choice of parental materials, crossing defensible manner. content through breeding are limited.
techniques, selection methods, and Research has shown that protein content
evaluation procedures) are likely to have is mainly influenced by environment
economic implications. To the extent that (and by genotype x environment
Assessing Potential
changes in the organization or operation interaction), so the genotype effect is
of a wheat breeding program affect the
Changes to a W heat generally very small (Bingham, 1979).
streams of costs and benefits, the Breeding Program Furthermore, given the known negative
economic outcomes that can be expected Under what circumstances might it be relationship between yield and protein
from the program will increase or advisable to incorporate physiology into content (O’Brien and Ronalds, 1984),
decrease. a wheat breeding program? In assessing any increases in protein obtained through
the organization and management of an selecting higher-protein cultivars are
Plant breeders are viewed by many,
existing breeding program and deciding likely to result in lower yields.
especially by those in the food
production and processing industries, as whether or not changes may be needed
to meet a particular objective, it will What level of breeding input
a resource that can be used to enhance
often be useful to review the following is appropriate?
the overall performance of the
preliminary questions before Once it has been decided that the
agricultural sector (Brennan, 1997).
undertaking formal economic analysis. research objective is best addressed
Precisely because they have this
through breeding, it is necessary to
capacity, plant breeders often face
Is the problem best determine what level of breeding input is
demands on their services that far exceed
addressed through appropriate. An appropriate level of
what they can realistically expect to
breeding? breeding input is one that can be justified
deliver. In a world of limited resources,
Before any changes are made to a in terms of the size of the expected
plant breeders therefore need some basis
breeding program, it is important to benefits. Generally these will be related
for deciding which among the many
determine whether the results being to the size of the target region: As the
demands being placed on them should
sought could be obtained more quickly size of the target region increases, so will
have priority. Although economic factors
and/or cheaply by some other means. the level of breeding effort that is
are often taken into account (explicitly
For example, if the research objective is justified. Brennan (1992)
or implicitly) when research priorities
78
and Maredia (1993) have shown that for attractive over the longer term. For On what basis will varieties
small target regions, it often will be example, Brennan and O’Brien (1991) be released for use by
appropriate only to select from among found that the incorporation of early- farmers?
breeding lines that have been imported generation, small-scale quality testing The procedures used to evaluate new
from elsewhere. But once the target into an Australian commercial wheat varieties prior to their release also merit
region increases beyond a certain size, it breeding program, while initially consideration, because evaluation
will be appropriate to establish a full- attractive, led to a lower economic return procedures can have important economic
fledged local crossing program. The for the breeding program (Box A). implications for breeding programs.
precise threshold values needed to justify
expansion of an existing breeding
program will depend, among other
things, on the characteristics of the target Box A: Incorporating Quality Selection into a
environment (area, average yield levels, W heat Breeding Program
use of improved varieties, etc.) and the
yield gains that can be expected by
A recurring question facing any wheat breeding program concerns when to begin selecting for quality
increasing the breeding input.
characteristics—for example, protein content or gluten level. Opinions differ as to what the appropriate
time is for starting quality testing. Some breeders feel that selecting for quality should be left until late in
What breeding strategy is the improvement process, after significant progress has been achieved in raising yield potential. Others
likely to be most efficient?
feel that selecting for quality in addition to yield should commence early in the breeding process, so that
Once it has been determined that the
low-quality materials are screened out early on.
problem is best handled through
breeding and that the size of the target Brennan and O’Brien (1991) used an economic framework to evaluate the efficiency of two alternative
region justifies a full-fledged crossing approaches to the problem. Their study focused on two Australian wheat breeding programs, one which
program, a breeding strategy must be performed early generation quality testing, and another which did not (see table). Both programs used the
decided. Wheat breeding can be pursued same amount of labor, and the number of crosses and lines sown in the F2 generation was identical. The
in many different ways. The initial two programs differed mainly in the stage at which quality testing was initiated, which caused different
choice of source materials is of course sets of lines to move through each program.
critical; if the source materials selected Small-scale tests for quality carried out early in the breeding process (F2 stage) were found to be less
for improvement contain a high expensive than tests carried out in later generations (F6 stage). This led to the prediction that early
proportion of favorable alleles for the generation testing would prove more cost effective. But when the economic returns of the program doing
problem being addressed, the chances of early generation quality testing were compared to those of the program in which quality testing was
success are greatly enhanced. After introduced at a later stage, they were found to be lower. The costs associated with the program that
choosing the source materials, the included early generation testing were slightly
breeder must determine how much effort higher, but the benefits were markedly lower over Economic returns to early versus
will be put into crossing, as compared to the longer term. The program without early late selection for grain quality in wheat.
selection and evaluation of the resulting generation quality testing was able to concentrate
lines. In addition to a wide range of so- exclusively on yield potential, enabling it to Program Program
called conventional breeding methods, A† B‡
achieve much more rapid rates of yield gains.
modern breeders also have the option of Admittedly, it also had a lower rate of quality Expected increase over
incorporating biotechnology techniques, increase, since less selection pressure was placed current varieties (%):
such as the use of genetic markers, tissue on quality, but when economic values were - yield 4.6 2.3
culture, etc. In deciding what types of assigned to yield levels and quality factors, the - quality 0.2 1.1
selection methods will be optimal, it is additional yield gains more than compensated for Total costs‡ (US$ 000) 353 369
important that the decision be driven by the lower levels of quality. The study thus showed Total benefits‡ (US$ 000) 3,710 2,557
what is best for the program and its that in the absence of a substantial premium for Benefit-cost ratio 10.5 6.9
outcomes, not simply by the availability quality, wheat producers and consumers will be † Quality-testing introduced in F for Program A, and F
of new tools or techniques (Brennan, disadvantaged if breeding programs opt to select 6 2
1997). Innovative technologies that for Program B.
for quality in the early generations at the ‡ Discounted to year of crossing at 5% per annum.
appear to be tremendously appealing in expense of yield improvement. Source: Derived from Brennan and O’Brien (1991).
the short run often turn out to be far less
3 The important issue of how many varieties should be released may also have to be decided by breeders, although usually this matter is left to some sort
of government-appointed varietal certification and release committee.
Figure 1. Undiscounted stream of costs and Figure 2. Effects of discounting the stream of Two alternative approaches can be used
benefits associated with a plant breeding costs and benefits associated with a plant to analyze marginal changes: 1)
program. breeding program. comparing only changes in costs and
Source: Figure 12 in Morris et al. (1992). Source: Figure 13 in Morris et al. (1992). benefits expected to result from the
addition of the physiology component
(partial budget analysis), or 2) comparing
the costs and benefits of the entire
breeding program with the physiology
component to the costs and benefits of
4 Depending on the degree to which farmers sell their products, and depending on the nature of the markets in which they sell their products, these
productivity gains may be transmitted in part or in whole to consumers.
Conclusions following these steps will not necessarily J.L. and Chung, O.K. (eds.). Proceedings,
International Wheat Quality Conference.
ensure that the “correct” decision will be pp. 363-76.
With funds for agricultural research taken. And as we have pointed out, the Brennan, J.P. (forthcoming). Efficiency in wheat
becoming increasingly scarce in most outcomes of research are by nature improvement research: A case study of wheat
countries, research administrators face improvement research in Australia. In: Maredia,
uncertain, so some of the parameters used M. and Byerlee, D. (eds.). Efficiency of
mounting pressure to ensure that available in the economic analysis will necessarily Investment in National and International Wheat
resources are used efficiently. Although be tenuous. But one big advantage of Improvement Research. CIMMYT Research
there can be no question that a properly invoking an economic framework of
Report. Mexico, D.F.: CIMMYT.
Brennan, J.P., and Khan, M.A. 1989. Costs of
organized and managed physiology analysis is that it forces decision makers Operating a Wheat Breeding Program. Rural
component has the potential to add value to think somewhat more systematically and Resource Economics Report No. 5.
to wheat breeding activities, this does not about the many factors that are likely to Division of Rural and Resource Economics.
necessarily mean that every wheat N.S.W. Agriculture and Fisheries, Sydney.
influence the outcome of investment Brennan, J.P., and O’Brien, L. 1991. An economic
breeding program should include one. decisions; this in turn increases the investigation of early-generation quality testing
likelihood of achieving a favorable in a wheat breeding program. Plant Breeding
This chapter has reviewed some basic 106(2):132-40.
concepts from investment analysis that outcome. Gittinger, J.P. 1982. Economic Analysis of
can be used to assess the desirability of Agricultural Projects. Second Edition. Johns
If decisions about the role of physiology Hopkins, Baltimore, Maryland.
investing in a physiology component of in wheat breeding are taken based partly Maredia, M.K. 1993. The economics of international
a breeding program. We have described a on economic considerations, then agricultural research spillovers: Implications for
series of steps that may be useful in changes made to the organization and
the efficient design of wheat improvement
research programs. Unpublished PhD
helping to formalize decisions that all management of existing breeding dissertation, Michigan State University, East
too often are still left to the “gut feeling” programs are likely to lead to genuine Lansing, MI.
of scientists and research administrators: improvements in efficiency. Morris, M.L., Clancy, C., and Lopez-Pereira, M.A.
1992. Maize research investment and impacts in
• establish that the problem is Improvements in efficiency in turn developing countries. Part I of 1991-92
appropriately addressed through enhance the flow of new varieties CIMMYT World Maize Facts and Trends: Maize
emanating from the breeding programs, Research Investment and Impacts in Developing
breeding;
Countries. Mexico, D.F.: CIMMYT.
leading to increases in farm-level O’Brien, L., and Ronalds, J.A. 1984. Yield and
• estimate in rough terms the costs and
productivity that will eventually benefit quality interrelationships amongst random
benefits of the current breeding F3 lines and their implications for wheat
program; both producers and consumers.
breeding. Austr. J. Agric. Res. 35(6):443-51.
Reynolds, M.P., S. Nagarajan, M.A. Razzaque, and
O.A.A. Ageeb (eds.). 1997. Using Canopy
Temperature Depression to Select for Yield
Potential of Wheat in Heat Stressed
Environments. Wheat Special Report No. 42.
Mexico, D.F.: CIMMYT.
FACTORS
87
C H APTER 7
Traits to Improve Yield in
Dry Environments
R.A. Richards, A.G. Condon, and G.J. Rebetzke1
Genetic increases in wheat yields in dry Breeding for specific physiological traits height. Genetic manipulation of
areas have not been as great as in more that are expected to impart a yield flowering time has been important to
favorable environments or where advantage in dry environments has been adjust the duration of vegetative growth,
irrigation is available. A likely reason notoriously difficult and unsuccessful. reproductive growth, and grain growth in
for this is that dry environments are Among the reasons for the lack of relation to water supply, frost, and
characterized by unpredictable and success is that considerable research evaporative demand. A reduction in plant
highly variable seasonal rainfall and, effort has been directed towards traits height has been universally important in
hence, highly variable yields. This that are unlikely to improve productivity, increasing the proportion of grain to
results in slow genetic advances in and traits that have a low heritability or biomass (harvest index), provided
breeding programs because genetic are difficult to measure. There may also biomass growth is not compromised.
variation in yield is masked by large be negative correlations between drought
The understanding of physiological and
genotype x year and/or genotype x adaptive traits and with yield. For
morphological traits that limit yield
location interactions. example, earlier flowering may result in
under drought has improved in recent
reduced biomass accumulation or may
It is interesting, but not surprising, that years, and this has opened up new
increase the risk of frost damage.
genetic increases in yield potential made opportunities. Before discussing these,
by selection in predictable irrigated Traits selected may also be inappropriate and how they may be used in a breeding
environments have resulted in broadly for the target region. For example, unless program, it is important, first, to
adapted wheats that are often well the trait conditioning survival at the establish how widespread drought may
suited to both favorable and low seedling stage also conditions response be and in which environments yield may
yielding, rainfed environments. This to drought at later stages, breeding for be limited by insufficient water. It is also
arises because genetic variation in traits survival during drought at the seedling important to establish, in any
that contribute to high yield in all stage may be totally inappropriate if environment, whether water is the
environments, such as a high harvest drought only occurs around flowering or primary factor limiting yields or whether
index, is greater in predictable grainfilling. Another reason any attempt other factors may override the water
environments and, therefore, more to breed for yield under drought is likely limitation.
likely to be selected under favorable to fail is that in many dry environments
conditions. There is no reason why low rainfall is not always the primary
genetic advance in favorable factor responsible for low yields; other
The Extent of Drought
environments should not continue to factors, such as soil mineral nutrition or
contribute to yield in less favorable soilborne diseases, may be the
and Its Nature
environments, provided germplasm is overriding constraints. Since it is often Globally, CIMMYT recognizes 12
widely evaluated in rainfed difficult to identify those other, less distinct mega-environments (MEs)
environments. However, a host of obvious factors, this may also limit where wheat is produced. Only three are
specific adaptations that may be genetic progress in dry environments. irrigated (Rajaram et al., 1995). Several
uniquely important in rainfed of the rainfed environments have high
Only two traits have had a major impact
environments can also be targeted to rainfall (>500 mm) but may experience
on improving yields in rainfed
achieve higher regional yields. intermittent drought or terminal drought
environments: flowering time and plant
in years when rainfall is below average.
1 CSIRO Plant Industry, Canberra, Australia.
88
Three MEs experience frequent droughts and mistakenly be attributed to drought. light and macro-nutrient conditions
and are considered marginal with respect There may be nutritional problems, probably indicate some disorder.
to food production. perhaps mineral toxicities due to soil pH;
Finally, a very effective way to identify
nutrients may be chronically low or even
There are subclassifications within each limiting factors is to grow a range of
too high; there may be soilborne
of the MEs. For example, ME4, one of probe genotypes, or other cereal species,
pathogens such as nematodes, root or
the largest rainfed environments, that are known to vary in their tolerance
crown fungal diseases such as take-all,
comprising about 33 million hectares, is or resistance to soil mineral disorders or
or rhizoctonia. All of these factors result
further subdivided into environments soilborne pathogens (Cooper and Fox,
in either poor root growth or diseased
with late drought (e.g., Mediterranean 1996). Relatively greater growth or yield
roots, and reduce water uptake, inducing
climates), early drought (e.g., Argentina) of one or more of the probe genotypes
symptoms of drought (Picture 1).
and regions where wheat is grown on may make it possible to diagnose a
residual moisture (e.g., parts of India There are a number of ways to determine particular problem.
following monsoon rains). whether soil-based factors limit yields
Another valuable way to determine
(Table 1). First, soil tests can be
Since drought can also be a problem in whether factors other than drought are
conducted to assess whether a range of
favorable environments, physiological more important in limiting yields is to
disorders such as pH or micro- and
means of minimizing drought stress or
macro-nutrient deficiencies or toxicities
improving water use efficiency may also
may limit yields. Second, roots can be Table 1. Methods to assess whether biotic
influence yield in high yielding, rainfed
examined around mid-tillering to or abiotic factors other than drought are
environments. Indeed, the best farmers limiting yields.†
determine the presence of take-all,
will often have the best yields as well as
rhizoctonia, or some nematode species
experience the most severe droughts Tests for soil pH or micro- and macro-nutrient
(e.g., cereal cyst nematode). Third, tiller deficiencies or toxicities
because their practices lead to the
development can be monitored. Tillering
greatest use of available soil water. Planting and subsequent growth of probe genotypes
in temperate cereals follows a very or species
Reducing the impact of drought may also
predictable pattern in favorable
be important in irrigated environments if Tiller development
conditions and therefore can be used to
it results in less water being used to
detect poor plant health. For example, Examination of roots
achieve high yield (i.e., high water use Soil moisture availability at harvest
primary tillers appear in the axils of
efficiency).
leaves 1, 2, 3 etc., and secondary tillers Calculation of water-use efficiency
appear in the axils of primary tiller † Methods are listed in the order in which they should be
leaves. Missing tillers under favorable
conducted during the crop cycle.
Is Drought the Primar y
Determinant of Yield in
a Dr y Environment?
Low rainfall is usually perceived to be
the most important factor resulting in low
yields in dry environments. However,
this may not always be true. Other
factors, such as disease, soil nutritional
problems, or even waterlogging at certain
times, may limit yields and should
probably be overcome as far as possible
before applying physiological
understanding to improve yields under
drought. Because foliar diseases are
easily observed, they are targeted in
breeding programs. However, other more
Picture 1. Droughted sections of fields caused by the root disease take-all and
insidious, not readily identifiable waterlogging during the vegetative period. Limiting the effects of these factors, principally
problems may also result in low yields through management, will increase yields in dry environments.
Table 3. Plant traits that may increase soil water use or root growth, or indicate deep rooting.
Universal or
Heritability Expected GxE environment-specific trait
Traits that increase soil water use
Deeper roots low high specific
Phenology high low specific
Seedling vigor high low specific
Tiller inhibition high low specific
Osmotic adjustment low high specific
Traits that indicate deeper roots
Canopy temperature intermediate high -
Stomatal conductance intermediate high - Picture 2. Greater seedling vigor, such as
Stay-green intermediate high - that achieved by the crop on the left, will
Leaf rolling intermediate high - reduce loss of soil water by evaporation
from the soil surface and limit weed growth.
Table 4. Traits that may improve plant establishment and early canopy development in wheat.
Universal or
Traits Heritability Expected GxE environment-specific trait
Highest priority
Long coleoptiles high low universal
Broad seedling leaves high low specific
Émbryo size high low specific
Specific leaf area intermediate high specific
Large coleoptile tiller intermediate high specific
Lower priority
Large grains high low universal
Fast emergence low low specific
Fast leaf expansion rate intermediate low specific
Low temperature
tolerance intermediate low specific
Crown depth intermediate intermediate specific
Picture 3. Semidwarf GA-sensitive wheats
Crown to shoot
with long coleoptiles establish better under
partitioning intermediate low universal
adverse conditions than GA-insensitive semid-
Leaf area ratio intermediate low specific
warf wheats with the Rht1 and Rht2 alleles.
Greater transpiration Table 5. Traits that can be selected to improve transpiration efficiency in wheat.
efficiency
Universal or
Transpiration efficiency (the ratio of dry Traits Heritability Expected GxE environment-specific trait
matter to transpiration), the other
component of water use efficiency, is Phenology high low specific
Seedling vigor high low specific
also amenable to improvement. There are
Carbon isotope discrimination high low universal
numerous ways to increase TE in wheat; Ash content ? high
the more important ones are given in NIR ? ?
Table 5. Perhaps the simplest way to Stomatal conductance intermediate high
improve TE is to ensure that the period of SPAD intermediate intermediate
SLA intermediate intermediate
maximum biomass increase occurs
Canopy temperature intermediate intermediate
during the coolest period. This capitalizes Glaucousness high low universal
on the fact that less water is required for Pubescence high low universal
growth when it is cool. To achieve this Residual transpiration ? high universal
may require a change in planting time so Leaf size and habit ? high universal
Natural soil salinity predates human Distribution of Areas affected by soil salinity are not
civilization. When early man, looking Saline Soils well defined, since detailed maps are
for better sources of livelihood, moved available for only a few. Consequently,
to arid lands along the riverbanks, he Salt-affected lands occur in practically global estimates vary widely (Flowers et
resorted to irrigated agriculture. With the all climatic regions, from the humid al., 1986). Of nearly 160 million hectares
practice of irrigation began salinity, the tropics to the polar regions. Saline soils of cultivated land under irrigation
first man-made environmental problem. can be found at different altitudes, from worldwide, about one-third is already
The earliest written account of salt lands below sea level (e.g., around the Dead affected by salt (Figure 1), which makes
dates back to 2400 BC and was recorded Sea) to mountains rising above 5000 salinity a major constraint to food
in the Tigris-Euphrates alluvial plains of meters, such as the Tibetan Plateau or production. It is the single largest soil
Iraq (Russel et al., 1965). The first time the Rocky Mountains. The occurrence of toxicity problem in tropical Asia
salt lands were associated with irrigation saline soils is not limited to desert (Greenland, 1984). In the wheat growing
was in northeastern Sumer, in the conditions; the problem has been areas of India, the combination of salt-
vicinity of modern Telloh. Salinity is reported in the tropical belts of Africa affected soils and poor quality
thought to have been partially and Latin America, and even in the polar groundwater (Figure 2) severely limits
responsible for the breakdown of the regions, particularly Antarctica. productivity.
ancient Sumarian civilization (Jacobson
and Adams, 1988).
1 Crop Improvement Division, Central Soil Salinity Research Institute, Karnal, 132 001 (Haryana), India.
101
Effect of Salinity /
Alkalinity on Plants
Crop species show a spectrum of
responses to salt, although all have their
growth and, eventually, their yield
reduced by salt. Salt effects are the
combined result of the complex
interaction among different
morphological, physiological, and
biochemical processes.
Salt-affected soils (Mha) Poor quality ground water (%)
Morphological effects
Gujarat Maharashtra Rajastham Madhya Pradesh
Morphological symptoms are indications
Haryana Punjab Bihar Uttar Pradesh of the injurious effects of salt stress. The
extent of inhibitory or adverse effects
Figure 2. Distribution of salt-affected soils and poor quality ground water in wheat-growing
can be known only by making critical
states of India.
comparisons with plants growing under
comparable conditions in normal soils.
A salt tolerance breeding program cannot • Localization of saline ions in old • Availability of suitable genetic
be successful in the absence of data on leaves so as to protect the relatively variability in the cultivated species or
the physiological mechanisms by which new and actively transpiring leaves their wild relatives.
plants cope with salinity stress. A precise (Yeo and Flowers, 1982). • Method of screening large numbers of
• Compartmentalization of ions within genotypes for salt tolerance.
understanding of the mechanism of
the leaf to avoid water deficit in the • A suitable breeding methodology.
expressed tolerance can help to resolve
leaf (Oertli, 1988).
the genetic base of the trait to
Picture 1. Screening for salt tolerant wheat genotypes in microplots under controlled salt
stress conditions.
Wheat is grown across a wide range of growth (about 20ºC), and there are resistant to freezing and may be injured
environments and is considered to have definitely differences in the growth rate at -1.8ºC (Single and Marcellos, 1974).
the broadest adaptation of all cereal crop of cultivars at low temperatures and, Differences in what is generally called
species (Briggle and Curtis, 1987). This consequently, in their adaptation to cool “frost tolerance” are less pronounced,
broad adaptation is due, to a large extent, climate. However, the term “cold although waxy or hairy lemma, palea,
to wheat’s cold tolerance, i.e. the ability tolerance” is most frequently used to and awns are thought to delay formation
to withstand temperatures much lower describe a plant’s response to freezing of ice in the tissue. Due to the limited
than 1-4ºC, considered the minimum temperatures, which have more dramatic genetic variation for frost tolerance,
temperature for growth (Figure 1). effects on the crop. Most often, freezing breeding efforts have been directed
temperatures affect autumn-sown wheat mostly to escaping frost by selecting for
In a general sense, cold tolerance in
during winter. Freezing tolerance refers later flowering.
wheat should refer to performance at
to the broader term of “winter hardiness,”
temperatures lower than the optimum for
an attribute of autumn-sown cereals that
is responsible for differences in “winter
survival” or “overwintering.”
Heat kill Winter survival is defined by Blum
Respiration accelerates; (1988) as “the final integrated plant
plants may lose weight
response to a multitude of stresses
Upper limit for photosynthesis
involved during and after freezing stress,
High temperature seed dormancy
including both external-physical and
Ideal for growth and development biotic stresses.” Even if plants are not
winter-killed, they can be affected by
Cold-hardening induced
freezing temperatures that may damage Picture 1. Differential winter damage in
the leaf, causing reduction in leaf area, head rows.
Spikelets and flowers freeze
Roots freeze
delayed growth, and plant debilitation.
Leaves freeze Considerable variation for winter
hardiness exists among cultivars, which
justifies dedicating extensive efforts to
Crowns of most winter-
hardy varieties killed this breeding objective (Pictures 1 and 2).
1 Research Institute for Cereals and Industrial Crops, Fundulea, 8264 Calarasi, Romania. Email: saulescu@valhalla.racai.ro.
2 Winter Wheat Breeding, CIMMYT/Turkey.
111
It should be noted that low, non-freezing (Andrews et al., 1974). Ice has high Gusta et al. (1997a) reported that the
temperatures (below 10ºC) at the critical thermal conductivity and can aggravate main factors responsible for winterkill in
stage of meiosis can also have dramatic the effect of low temperatures. It also the Great Plains of North America are
effects on wheat by causing male-sterility has low gas permeability and may, in long periods of cold-induced desiccation,
and, consequently, low yields. Genetic extreme cases, smother or suffocate poor acclimation conditions in autumn,
differences in the response to this stress plants by depriving them of oxygen and unpredictable timing and duration of
are known to exist (Qian et al., 1986, (Poltarev et al., 1992). extremely cold temperatures, whereas the
Saulescu et al., 1997) but, because of its primary cause of winterkill in western
Finally, low temperatures or snow can
relatively rare occurrence, little effort is Canada is freeze-induced desiccation. For
cause indirect damage through:
directed towards breeding for tolerance eastern North America, Olien (1967)
other than selecting for an appropriate • frost heaving due to the formation of found that winterkill is most likely to
flowering time that allows plants to ice in the soil. The ice pushes the occur during low temperature stress
escape the stress. plants upward, breaking and exposing following a midwinter thaw, when the
the roots; crown tissues have high moisture content.
• snow mold, caused by fungi in areas Correct evaluation of the frequency with
with long-lasting snow cover. The which these or other factors can affect
Stress Factors Involved most damaging fungus affecting
winter survival in the target area is
in W interkill winter survival is pink snow mold
essential for making a better choice of
(Microdochium nivale (Fries) Samuel
The reasons for winterkill in wheat, as and Hallet), previously known as parents and testing procedures in a
well as the extent of the damage, vary Fusarium nivale (Fr) Ces. (Hömmö, breeding program; this can also improve
greatly from region to region and from 1994). Although Microdochium resource allocation efficiency.
year to year. The main factors causing nivale cannot survive freezing, it is
Wheat plants can cope with each of the
winterkill (alone or in combination) are tolerant to low temperatures and
above mentioned winter stress factors
related to low temperature per se (such as severely damages plants in the 0-5ºC
temperature range. Other, less through different genetic and
extreme air or soil temperatures, below
important fungi causing snow mold physiological mechanisms. For example,
the critical temperature of a particular
are Typhula spp., the pathogen for a plant’s freezing tolerance and snow
wheat cultivar):
speckled snow mold or typhula mold resistance are based on different
• inadequate hardening, due to late blight, and Sclerotinia borealis, genetic mechanisms (Hömmö, 1994).
emergence in autumn or a sudden which causes sclerotinia snow mold. However, the basic process behind most
drop in temperature; events leading to winterkill is freezing, or
The relative importance of stress factors
• long periods of cold-induced formation of ice in plant tissues (Figure 2).
desiccation (Gusta et al., 1997a); causing winterkill can vary greatly
Freezing damage is in general not a
• prolonged periods of low sub-zero among regions. In the Ukraine, an
consequence of low temperature per se,
temperatures; in particular, mid-winter analysis of data from the last 100 years
but rather the result of cellular
temperatures below -15ºC result in the showed that winterkill was caused by
dehydration brought about by extra-
rapid loss of winter hardiness (Gusta low temperatures in 35% of cases, by
et al., 1997b); cellular ice crystallization. Cellular
alternate freezing and thawing in 26% of
• alternate freezing and thawing, which membranes have been recognized as the
cases, and by ice encasement in 22% of
causes increased injury from ice primary sites of freezing injury (Hincha
years when significant winter damage
crystal growth with each freeze and Schmitt, 1994).
occurred (Poltarev et al., 1992).
(Olien, 1969). Wisniewski et al. (1997) stated that the Freezing tolerance is defined as the
Another factor responsible for winterkill critical factors that affect winter survival ability of plants to survive ice formation
is ice encasement, a major cause of plant in Poland are low temperature, freeze- in extracellular tissues without significant
death in areas of high rainfall and induced desiccation, and infection by damage to membranes or other cell
fluctuating temperatures during winter pathogenic fungi. components.3 It is the result of
physiological, chemical, and physical
3 For the sake of clarification, it should be noted that intracellular ice formation is always lethal. The chemical potential in the intracellular solution
must be equal to the chemical potential of the external solution or the ice. This equilibrium is attained through removal of intracellular water. To avoid
cellular dehydration under freezing stress, the osmotic potential of intracelluar solutions is increased as the osmotic potential of extracellular solution
decreases. For a detailed discussion of the physiological processes during freezing stress, see Blum (1988).
4 Cereals differ greatly in their ability to survive low temperatures. The most cold tolerant rye cultivars are killed at around –34oC, wheat cultivars at
around –23oC, and barley at around –18oC.
5 Ryabchun et al. (1995) recommend adding 36 h at -5ºC, 56 h at -7ºC, 24 h at -9ºC, and 14 h at -10ºC of artificial hardening to the level achieved through
natural hardening in the field by 14-25 November under conditions in Kharkov, Ukraine.
Wheat is the most widely grown cereal affect the crop. Perhaps the greatest When consulted, representatives of
in temperate environments, and is also challenge to understanding the national agricultural research systems
cultivated in many tropical cropping physiological problems associated with (NARSs) from the major wheat-growing
systems, where it is often grown as the heat stress is to encompass the diversity regions in the developing world
winter season crop in rotation with a of hot environments all over the world identified heat stress as one of their top
number of other crops—for example, (Figure 2). Continual heat stress affects research priorities (CIMMYT, 1995).
with maize in Africa, rice in Asia, and approximately 7 million ha of wheat in
soybean in Latin America (Figure 1). developing countries, while terminal
Among the numerous advantages of heat stress is a problem in 40% of
cultivating wheat in this niche are that it temperate environments, which cover CIM MYT / NARS
is stress tolerant, relatively high 36 million ha. Continual heat stress is Collaboration on Heat
yielding, and easy to transport and defined by a mean daily temperature of Tolerance
store. over 17.5°C in the coolest month of the
season (Fischer and Byerlee, 1991), and Breeding efforts by a number of national
There are also disadvantages linked to over 50 countries (importing more than wheat breeding programs has resulted in
growing wheat in tropical areas; 20 million tons of wheat per year) the release of germplasm adapted to
foremost among them are the different experience this type of stress throughout warm growing environments, such as in
types of high temperature stress that the wheat cycle. Egypt and Sudan (AbdElShafi and
Ageeb, 1994), India (Tandon, 1994),
124
Bangladesh (Razzaque et al., 1994), and The information emanating from the Physiological Traits
Uruguay (Pedretti and Kohli, 1991). IHSGE may be useful in establishing Associated with Heat
CIMMYT has been actively involved in indirect selection criteria for heat
Tolerance
many of these regions (Kohli et al., tolerance. The application of some of
1991; Ortiz-Ferrara et al., 1994). these traits to breeding will be Genetic diversity for heat tolerance in
Collaboration between CIMMYT and discussed in subsequent sections, but cultivated wheat is well established
NARS on physiological aspects of heat first we will present a brief review of (Midmore et al., 1984; Rawson, 1986;
tolerance in wheat started in 1990, with some of the physiological traits Wardlaw et al., 1989; Al-Khatib and
the establishment of a network involving associated with heat stress. Paulsen, 1990; Reynolds et al., 1994).
wheat scientists in Bangladesh, Brazil, Photo-assimilation is more likely to be
Egypt, India, Nigeria, Sudan, and yield-limiting under heat stress than in
Thailand. Table 1. Genetic correlations between temperate environments, especially as
morphological traits and wheat yields for 10 stress typically intensifies during
Collaborative experiments conducted by
varieties averaged over 16 low relative humidity grainfilling, when demand for assimilates
network scientists were named the
environments in ME5, IHSGE 1990-94. is greatest. This is borne out by the
International Heat Stress Genotype
observation that under stress, total above-
Experiment (IHSGE) (Reynolds et al.,
1992; 1994; 1997; 1998; Reynolds, Trait Genetic correlation ground biomass typically shows a
stronger association with yield than with
1994). The IHSGE was grown in wheat-
Final biomass (above ground) 0.88** partitioning, i.e., harvest index (Table 1);
growing areas classified by CIMMYT as
Grains/m2 0.77** the situation is usually reversed under
heat stressed, i.e., CIMMYT mega-
Grains/spike 0.67* temperate conditions.
environment 5 (ME5). The main
Harvest index 0.51
objectives of the IHSGE were to Hence traits affecting radiation use
Kernel weight -0.10
establish the degree of genotype by efficiency (such as early ground cover,
Spikes/m2 0.0
environment interaction (GxE) in ME5, stay-green, and photosynthetic rate) could
evaluate potential physiological Days to anthesis 0.83** be expected to be important under heat
screening techniques by observing Days to maturity 0.81** stress. Although early ground cover
genetic diversity for traits and their Plant height 0.20 seems to be important in an agronomic
association with heat tolerance, and context (Rawson, 1988; Badaruddin et
improve our understanding of the % ground cover (anthesis) 0.67* al., 1999), variation in this trait among
physiological and genetic basis of heat Biomass at anthesis 0.35 genotypes does not seem to be associated
tolerance. Plant dry weight (5-leaf stage) -0.45 with heat tolerance (Table 1). The stay-
% ground cover (5-leaf stage) -0.30 green trait has been used widely in
There were three main outcomes of the Plants/m2 -0.15 breeding for heat tolerance, partly as an
study. First, cluster analyses of over 40
indicator of disease resistance (Kohli et
hot sitex year combinations indicated * Denotes significance at ≤ 0.05, ** significance at ≤ 0.01.
al., 1991). Physiological evidence
that most interaction between sites and
indicates that loss of chlorophyll during
genotypes was accounted for by relative
grainfilling is associated with reduced
humidity (RH). Hence low RH sites
yield in the field (Reynolds et al., 1994).
(e.g., Sudan, Mexico, and India) and
Studies in controlled environments have
high RH sites (e.g., Bangladesh and
revealed genetic variability in
Brazil) showed less Gx E within RH
Table 2. Genetic correlations (Rg) for photosynthetic rate among wheat
groups than when comparison was done
physiological parameters measured in cultivars when exposed to high
across RH groups (Reynolds et al., 1998;
Tlaltizapan, Mexico, and wheat yields for 10 temperatures (Wardlaw et al., 1980;
Vargas et al., 1998). This kind of
varieties averaged over 16 low relative Blum, 1986).
analysis indicates that breeding for these
humidity environments, IHSGE 1990-94.
two broad environments should be Such differences in photosynthesis under
undertaken as separate objectives. Physiological trait R(g) heat stress have been shown to be
Second, data collected on IHSGE lines associated with a loss of chlorophyll and
in the low RH sites showed consistent Canopy temperature depression 0.86** a change in the a:b chlorophyll ratio due
association between yield and a number Membrane thermostability 0.81** to premature leaf senescence (Al-Khatib
of morphological traits (Table 1). Third, Leaf chlorophyll (grainfilling) 0.72** and Paulsen, 1984; Harding et al., 1990).
physiological data collected in Mexico Leaf conductance (heading) 0.63* Studies at CIMMYT comparing 16
showed that several parameters were Photosynthesis (heading) 0.63* diverse semi-dwarf wheats demonstrated
associated with performance at genetic variability for photosynthetic rate
international low RH sites (Table 2). * Denotes significance at ≤ 0.05, ** significance at ≤ 0.01. under heat-stressed field conditions
T
O-1
CHG
EMISSIVITY
2:3
randomly derived sister lines using a
number of relevant crosses and
heritability of traits established, as
outlined in the introduction of this book.
Evidence for applying three traits
(namely, canopy temperature
depression, leaf conductance, and
membrane thermostability) in selecting
for heat tolerance is presented in the
following sections; sufficient evidence
has been collected on these traits to
suggest their potential as breeding tools.
Figure 3. Potential use of canopy temperature depression in a breeding program.
Nonetheless, if these techniques have
not been evaluated in a given breeding
environment, they should first be
evaluated, as outlined in chapter one, Table 3. Summary of heat stress mechanisms previously reported for wheat and their
before being applied to mainstream association with yield in the IHSGE.
breeding operations. Reported heat stress mechanism Accounting for genetic variation in yield in IHSGE
Canopy temperature Accelerated development Yes, lateness associated with higher yield in many
depression (Midmore et al., 1984) environments
As discussed earlier, experimental data Stand establishment (Rawson, 1988) No, poor correlation with early growth
have shown a clear association of CTD
with yield in both warm and temperate Evaporative cooling (Idso et al., 1984) Yes, strong correlation of CTD with yield
environments. CTD shows high genetic
Inhibition of meiosis No, sterility not observed. Grain:spikelet ratio not
correlation with yield and high values
(Saini et al., 1983; Zeng et al., 1985) correlated with yield
of proportion of direct response to
selection (Reynolds et al., 1998), Sensitive growth phase Partial least squares analysis confirmed spike growth
indicating that the trait is heritable and (Fischer, 1985; sensitivity, especially to high night temperatures (Vargas
therefore amenable to early generation Shpiler and Blum, 1991) et al., 1988)
selection. Since an integrated CTD
value can be measured almost Photosynthesis/chlorosis Yes, high association of photosynthesis and stay-green
instantaneously on scores of plants in a (Al-Khatib and Paulsen, 1990; with yield in field plots
small breeding plot (thus reducing error Shpiler and Blum, 1991)
normally associated with traits Thylakoid thermostability Preliminary data on IHSGE lines confirms association of
measured on individual plants), work (Moffatt et al., 1990) chlorophyll fluorescence with yield (Balota et al., 1996)
has been conducted to evaluate its
potential as an indirect selection Membrane thermostability (MT) Yes, MT measured on seedlings and flag leaves associated
criterion for genetic gains in yield. CTD (Shanahan et al., 1990) with yield at several sites
is affected by many physiological Inhibition of starch synthase No clear evidence, but yield not associated with TGW
factors, which makes it a powerful (Bhullar and Jenner, 1986; Rijven, 1986)
4
When crop is sown in
rows, point thermometer
at an angle to rows, not 3
parallel to them.
IRT
1
6.0 7.0 8.0 9.0
Canopy temperature depression (C)
At low leaf area
index, point IRT at Figure 7. Regression of yield on CTD
oblique, NOT acute, measured after heading for 40 recombinant
angles to avoid inbred lines from a cross between lines
intercepting soil. contrasting in heat tolerance (Seri 82* Siete
Cerros 66), Tlaltizapan, Mexico, 1995-96.
Source: Reynolds et al. (1998).
Figure 5. How to view a plot to avoid including soil temperature when measuring canopy
temperature depression with an infrared thermometer (IRT).
Grain yield (t ha-1) Table 5. Correlation coefficients between yield, averaged over two cycles at six locations of
5.5 the IHSGE (1990-92), and CTD of 16 wheat lines measured at different stages of
development, December and February sowings, Tlaltizapan, Mexico, 1992-93.
5.0
4.5 CTD December CTD February
4.0 Location Pre-anthesis Anthesis Post-anthesis Pre-anthesis Anthesis Post-anthesis
3.5
Brazil 0.45 0.60* 0.50* 0.68** 0.52* 0.68**
3.0 Egypt 0.73** 0.91** 0.91* 0.82** 0.79** 0.78**
2.5 India 0.33 0.56* 0.62** 0.60** 0.37 0.64**
2.0 Sudan 0.71** 0.91** 0.88** 0.77** 0.75** 0.71**
Tlaltizapan 0.66** 0.84** 0.78** 0.50* 0.53* 0.43
1.5
5.0 6.0 7.0 8.0 9.0 Average 0.58 0.76 0.74 0.67 0.59 0.65
Canopy temperature depression (C) correlation
Figure 6. Relationship of mean grain yield * Denotes significance at ≤ 0.05, ** significance at ≤ 0.01.
to mean CTD for 23 genotypes, averaged Source: Reynolds et al. 1994.
over two sowings, Tlaltizapan, Mexico,
1992-93.
Source: Amani et al. (1996).
More than one third of the world’s The effects of waterlogging are most Waterlogging can affect other irrigated
irrigated areas suffer occasional or more widespread in the irrigated rice-wheat areas in Asia besides the rice-wheat
frequent waterlogging (Donmann and regions of South and Southeast Asia growing regions. Wheat-producing
Houston, 1967). Waterlogging has been (i.e., China, Vietnam, Thailand, areas in Egypt, Sudan, and Nigeria also
shown to limit wheat yields in many Bangladesh, Nepal, India, and Pakistan) suffer regularly from waterlogging. In
regions of the world; an area estimated and in the southern United States parts of Africa and Latin America,
at 10 million ha is waterlogged each (i.e., Georgia, Mississippi, and heavy rainfall combined with heavy
year in developing countries (Sayre et Louisiana). A common denominator in clay soils creates waterlogging that
al., 1994). Waterlogging occurs when these countries is that rice rotations are limits wheat production. In the
rainfall or irrigation water collects on practiced on much of the land. Soils are traditional wheat-growing regions in
the soil surface for prolonged periods generally puddled to restrict water the Ethiopian highlands, downpours are
without infiltrating the soil. Soil percolation and create flooded heavy and prolonged during the rainy
characteristics that contribute to conditions for rice cultivation. Due to season. Hence waterlogging is a
waterlogging include soil physical soil puddling, wheat that follows rice in common occurrence at the beginning of
properties that allow formation of a the drier season is planted under less the wheat cycle. The situation is further
crust on the soil surface or of a pan in than optimal soil physical conditions. exacerbated by the black vertisols in
the subsoil. Waterlogging can also occur The soil pan that was created Ethiopia, consisting of heavy clays that
when the amount of water added intentionally for rice cultivation is often inhibit infiltration, swell, and crack
through rainfall or irrigation is more left undisturbed and may create a severely. Waterlogging limits wheat
than what can percolate into the soil barrier for water movement, causing yields in Australia due to rising
within one or two days. waterlogging when excessive irrigation groundwater (Grieve et al., 1986;
or rainfall occurs. McDonald and Gardner, 1987; Meyer
Waterlogging occurs in many wheat
and Barrs, 1988).
growing regions around the world, In South Asia, wheat is a relatively new
especially irrigated and high rainfall option within rice rotation schemes.
environments. In irrigated regions, the Some farmers, accustomed to applying
main culprit seems to be the lack of generous amounts of water to rice, tend Conditions and
proper drainage systems. Irrigation to over-irrigate their wheat crop. Symptoms Associated
facilities do not allow easy drainage of Additionally, many rice-wheat soils are
with Waterlogging
excess water, sometimes due to poorly silt or loam and susceptible to crusting,
kept irrigation canals from which water which creates waterlogging by Except at sowing or during early
seeps out. Major examples are the Indian restricting percolation from the surface. germination, waterlogging will not
Subcontinent, certain river basins in Declines in organic matter in the topsoil generally destroy wheat plants nor
China, and the Nile River Delta in Egypt. of South Asia are well documented and affect plant establishment (Musgrave,
In the northern Indo-Gangetic Plains of also contribute to poor soil physical 1994). The major morphological and
India alone, 2.5 million ha of wheat are quality (FAO, 1994; Hobbs and Morris, biochemical effects will be discussed in
affected by irregular waterlogging 1996; Nagarajan, 1998). detail later, but under mild
(Sharma and Swarup, 1988). waterlogging wheat plant growth is
1 Principal Scientific Officer, Wheat Research Centre, Bangladesh Agriculture Research Institute, Joydebpur, Gazipur, Bangladesh (same mailing
address as CIMMYT below).
2 CIMMYT Natural Resources Group, P.O. Box 6057, Gulshan, Dhaka-1212 Bangladesh.
3 Senior Scientific Officer, Wheat Research Centre, Bangladesh Agriculture Research Institute, Joydebpur, Gazipur, Bangladesh (same mailing address
as CIMMYT above).
4 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
136
usually stunted, bottom leaves senesce, (Belford et al., 1985). Generally, the root respiration, though on a more
tiller survival is reduced, and florets may wheat plant’s tolerance to waterlogging limited basis than in aerobic conditions.
become sterile. increases as it ages, and the detrimental The process is accelerated if temperatures
effect on yield decreases (Meyer and are elevated. Genetic variability for this
High temperatures tend to exacerbate the Barrs, 1988). Once the wheat crop is trait has been documented in the literature
negative effects of waterlogging. When established, many genotypes can (Cao et al., 1995).
aerobic soil conditions re-occur, plant withstand waterlogging up to 10 days
growth resumes slowly. Consequently, with no yield loss, if the wheat leaves Winter wheat areas may also be prone
wheat yields are affected. are not submerged. Wheat crops can to waterlogging. Winter wheats are
make an amazing recovery following sometimes grazed and allowed to re-
An entire field will rarely be grow for grain production in the spring.
waterlogged; waterlogging is usually early waterlogging stress, if supplied
with extra nitrogen. Trampling of saturated pasture soils by
restricted to the lower lying areas of a cattle can cause restricted water
field (Picture 1). Waterlogging occurs In waterlogged soils and in the roots of movement and waterlogging.
when the soil is fully saturated, and plants growing in them, exceptionally
standing water replaces the air in the soil high levels of ethylene may build up, The literature documents some
pore spaces. There is a lack of oxygen in given that ethylene diffuses more slowly tolerance of winter wheats to
the soil, restricting aerobic respiration by in water than in aerated soil. The first waterlogging (Musgrave, 1994). Yet this
growing roots and other living response of a wheat plant to anaerobic may not be true tolerance, since the
organisms. Soil chemical properties conditions involves its biochemical colder soil temperatures associated with
change when anaerobic conditions pathways as a response to the lack of waterlogging in winter-wheat-growing
persist for several days, increasing the respiration by root cells. Various areas reduce the amount of oxygen
availability of some major or minor hormones are stimulated and required for root respiration. Thus yield
elements while decreasing the transported to the leaves, causing early reductions associated with waterlogging
availability of others. Plant transpiration senescence of older leaves within days in colder areas are not as great as those
is affected until wheat roots recover (Dong et al., 1983; Dong and Yu, 1984). in the more temperate and tropical areas
(when soil aerobic conditions recur) or Seminal roots are generally killed or of the world. On the other hand, some
adapt to the anaerobic conditions. their growth greatly restricted (Huang studies show soil oxygen decline under
However, extended waterlogging will and Johnson, 1995). waterlogging is rapid at most
result in root death. Waterlogging also temperature ranges (Trought and Drew,
limits the wheat plant’s nutrient uptake However, some wheat genotypes have 1982). It should also be noted that
by reducing plant transpiration and nodal or adventitious roots that begin winter wheats are longer maturing and
diminishing root function. aerenchyma cell formation. hence less sensitive to waterlogging
Aerenchyma is tissue that can carry than the earlier maturing spring wheats
Another effect of waterlogging is to oxygen from the leaves to the roots (Gardner and Flood, 1993).
stimulate the production of certain plant under anaerobic conditions to maintain
hormones. In anaerobic conditions these
hormones are released from the roots in
greater concentrations and may affect
leaf and root responses. Ethylene is
produced both by the roots and by
microorganisms in waterlogged soils.
The hormonal effects of ethylene
released under waterlogging are
attracting a great deal of interest. Water
acts as a barrier to the escape of ethylene
produced in roots and other submerged
tissue. Ethylene is known to be a trigger
(not a promoter) of leaf senescence
(Dong et al., 1983).
Waterlogging during sowing or
germination generally kills the seed or
seedling. The seedling’s radicle and
roots do not adapt readily to
waterlogging or are more susceptible to
seedling diseases that may follow Picture 1. Non-uniform waterlogging in a wheat field in Bangladesh.
Picture 3. Waterlogging
reduces the number of
Picture 2. Lower leaf chlorosis. spike-bearing tillers.
Realistic but cautiously optimistic Figure 2. Waterlogging at different growth Figure 4. Waterlogging at different growth
conclusions can be drawn based on the stages (days after sowing-DAS) affected stages (days after sowing-DAS) affected
above review of the literature and on grains per unit area in 1995-96 at grain yield in 1995-96 at Joydebpur,
data from the case study in Bangladesh. Joydebpur, Bangladesh. Bangladesh.
Rainfall during or just prior to harvest Extent of the Problem Damage Caused
can cause wheat grain to germinate
while still on the spike (Figure 1). This Rainfall can cause extensive PHS As its name indicates, preharvest sprouting
phenomenon, called preharvest damage in most wheat-growing regions; begins before the grain is harvested, while
sprouting (PHS), reduces yield, lowers however, some areas are more prone it still on the spike. The process is set in
test weight, and adversely affects the than others to its occurrence. Although motion by rainfall, during which the seed
milling and baking quality of harvested many of these regions are found in imbibes water. This causes germination to
grain. Farmers receive lower prices for developed countries, significant areas of begin as starch reserves in the grain
sprouted grain and, in severe cases, the developing world are also affected. endosperm are hydrolized through the
their harvests may be downgraded to Northern Europe, the Pacific Northwest action of germinative enzymes called
animal feed. The occurrence of PHS is of the United States, and the wheat amylases. The embryo swells and grows as
generally erratic and as difficult to growing areas of central Canada and it consumes the hydrolized carbohydrate
predict as rainfall in most wheat- northeastern Australia periodically reserve.
growing areas. Researchers have, suffer PHS damage. In developing
countries the Southern Cone of South The test weight and flour milling yield of
however, been able to provide farmers sprouted grain are considerably lower
in areas prone to PHS with a degree of America, encompassing parts of Chile,
Argentina, and Brazil, and the wheat- than those of non-sprouted wheat. Bread
protection. This chapter attempts to produced from sprouted grain has poor
outline the role of physiology and growing areas of eastern Africa are
prone to PHS. The damage is more loaf volume and crumb structure, and is
plant breeding in the wider effort to unsuitable for marketing (Figure 2). The
develop strategies to combat this pronounced in regions where white-
grained wheat is grown. Red-grained quality of flat breads and chapatis is less
intractable problem. affected by the use of sprouted grain, but
wheat is more tolerant to the problem,
since there is an association between their texture is nevertheless impaired,
grain color and grain resulting in a less favorable product.
dormancy, the primary Sprouted grain causes discoloration of
mechanism of PHS Chinese noodles and spaghetti, lowering
tolerance (Gale, 1989). the value of these products as well.
Figure 1. Two spikes damaged by pre-harvest Figure 2. Bread produced from sprouted grain (left) and from sound grain (right).
sprouting (left) and a sound spike (right).
1 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico 06600.
145
Tolerance Mechanisms Screening Methods and buy, and many scientists cannot afford
4HERE ARE NO REPORTS OF DIRECT YIELD 'ENOTYPE AND 9IELD #ROP PHENOLOGY
RESPONSE TO EITHER HIGH WIND OR VAPOR 0OTENTIAL 4HE ABOVE COMMENTS ON THE IDENTIlCATION
PRESSURE DElCIT BUT NEGATIVE RESPONSES OF OPTIMUM mOWERING DATES
HAVE BEEN SUGGESTED !TMOSPHERIC 'ENETIC VARIATION FOR 9P AT A PARTICULAR NOTWITHSTANDING THE RETROSPECTIVE
POLLUTIONOZONE IN PARTICULARCAN LOCATION AND ESPECIALLY HISTORIC PROGRESS APPROACH HAS SOMETIMES REVEALED SHIFTS
REDUCE YIELD !NOXIA ASSOCIATED WITH IN 9P MUST REmECT GENETIC VARIATION IN CROP PHENOLOGY ASSOCIATED WITH YIELD
TRANSIENT WATERLOGGING AND PERHAPS FOR THE EFlCIENCY WITH WHICH THE IMPROVEMENT )N THE CASE OF WHEAT
mOOD IRRIGATION AND HIGH MECHANICAL CROP INTERACTS WITH THE EXTERNAL YIELD
MODERN VARIETIES HAVE TENDED TO HAVE
IMPEDANCE COULD HAVE SUBTLE NEGATIVE CONTROLLING FACTORS DISCUSSED ABOVE A REDUCED GERMINATION
TO
ANTHESIS PERIOD
EFFECTS ON YIELD DESPITE APPARENT OPTIMAL #URIOUSLY HOWEVER SUCH REASONING HAS !$ IN DAYS ALTHOUGH SOME RECENT
SOIL MANAGEMENT NOT USUALLY FACILITATED THE IDENTIlCATION OF #)--94 CULTIVARS APPEAR TO BE
GENOTYPIC TRAITS ASSOCIATED WITH HIGH 9P REVERSING THIS TREND /F COURSE A SHORTER
!LL THESE EXTERNAL FACTORS AFFECTING 4RADITIONALLY CROP SCIENTISTS HAVE SOUGHT GERMINATION
TO
ANTHESIS PERIOD MAY BE
9P VARY SUBSTANTIALLY AND TO SOME TO UNDERSTAND YIELD PROGRESS THROUGH THE DESIRABLE FROM A CROPPING SYSTEMS
EXTENT INDEPENDENTLY IN THE WORLDS TRAIT CHANGES SEEN IN HISTORIC SETS OF PERSPECTIVE EVEN IF 9P DOES NOT INCREASE
WHEAT
GROWING LOCATIONS REPRESENTING CULTIVARS EXHIBITING YIELD PROGRESS DUE 6ARIATION AMONG THE PHENOLOGICAL SUB
COMBINATIONS OF LATITUDE LONGITUDE AND TO BREEDING A SO
CALLED RETROSPECTIVE PERIODS WITHIN THE GERMINATION
TO
ALTITUDE 4HEY CAN ALSO VARY AT THE SAME APPROACH &ROM THIS POINT OF VIEW ANTHESIS PERIOD HAS NOT BEEN STUDIED IN
LOCATION BECAUSE OF YEAR AND SOWING &EIL AND 3LAFER ET AL THIS CONTEXT BUT COULD BE A SIGNIlCANT
DATE EFFECTS %XPERIENCE OVER MANY YEARS HAVE THOROUGHLY REVIEWED GENETIC DETERMINANT OF 9P &ISCHER
HAS USUALLY LED TO IDENTIlCATION AT IMPROVEMENT OF YIELD POTENTIAL OF SMALL $URUM WHEATS AND TRITICALES APPEAR TO
EACH LOCATION OF THE OPTIMUM mOWERING GRAIN CEREALS AND WHEAT RESPECTIVELY HAVE A LONGER POST
ANTHESIS PERIOD AND
OR ANTHESIS DATE AND ACCORDINGLY THE ! SECOND SOMEWHAT DISTINCT APPROACH SOME CALCULATIONS OF 9P SUGGEST THAT
OPTIMUM SOWING DATE .EVERTHELESS THE HAS BEEN TO PREDICT FROM PHYSIOLOGICAL IN BREAD WHEAT YIELD CANNOT INCREASE
CLIMATE VARIES SUFlCIENTLY THAT 9P WITH UNDERSTANDING THE IDEAL PLANT TYPE FOR SUBSTANTIALLY WITHOUT AN EXTENSION OF THIS
THE CURRENT BEST
ADAPTED CULTIVAR PLANTED MAXIMUM YIELD AND THEN TO CONSTRUCT PERIOD 0HASIC DEVELOPMENT IS DISCUSSED
ON THE OPTIMUM SOWING DATE RANGES FROM THIS PLANT IN ORDER TO TEST AND HOPEFULLY IN MORE DETAIL IN THE CHAPTER BY 3LAFER
TO THA ACROSS LOCATIONS AND AT VALIDATE THE PHYSIOLOGICAL HYPOTHESIS
LEAST ACROSS ANY
YEAR PERIOD AT $ONALD ! VARIATION ON THIS 2EDUCED STATURE AND HARVEST INDEX
ANY LOCATION !S A RULE OF THUMB 9P IN APPROACH IDENTIlES SINGLE TRAITS OR -ANY STUDIES HAVE SHOWN THAT IN
THA AT MOISTURE WITH TODAYS BEST IDEOTRAITS FOR IMPROVEMENT WHICH ARE WHEAT THE USE OF MAJOR DWARlNG GENES
CULTIVARS IS GIVEN BY AMENABLE TO TESTING THROUGH THE MOST NOTABLY THOSE FROM .ORIN
HAS
DEVELOPMENT OF NEAR
ISOGENIC LINES BROUGHT ABOUT SIGNIlCANT AND LARGELY
9P 041 n 4 n 041
4
$ONALDS IDEOTYPE APPROACH WAS UNANTICIPATED INCREASES IN 9P QUITE
)N EQUATION 041 IS THE MEAN DAILY REVIEWED BY 3EDGLEY AND -ARSHALL INDEPENDENTLY OF THE INCREASE IN LODGING
SOLAR RADIATION -* M
DIVIDED BY WHILE !USTIN GIVES A MORE RESISTANCE 4HIS IS REmECTED IN AN INCREASE
THE MEAN DAILY TEMPERATURE IN # MINUS RECENT PHYSIOLOGICAL VIEW OF DESIGNING IN HARVEST INDEX () OF lNAL GRAIN
# DURING THE
DAY PERIOD UP TO THE CROPS FOR HIGHER YIELD 4HE RESULTS OF BOTH BIOMASS BUT NO CHANGE IN lNAL BIOMASS
END OF ANTHESIS USUALLY BETWEEN AND RETROSPECTIVE AND IDEOTYPE STUDIES WILL )N WHEAT REDUCED HEIGHT DUE TO MINOR
-* M
#
AND 4 IS THE MEAN THEREFORE BE REVIEWED ONLY BRIEmY HERE GENES ALSO APPEARS TO INCREASE HARVEST
DAILY TEMPERATURE DURING GRAINlLLING 4HE READER IS DIRECTED TO 2EYNOLDS ET AL INDEX AND 9P A PROCESS THAT HAS BEEN
USUALLY
# 4HIS IS DERIVED FROM FOR ADDITIONAL AND RECENT EXPLOITED BY BREEDERS THROUGHOUT THIS
RELATIONSHIPS IN 3AYRE ET AL DETAIL ON 9P IN WHEAT AND TO #ASSMAN CENTURY
FOR A USEFUL CONSIDERATION OF THE (OWEVER IN THE LAST YEARS WE SEEM
PARALLEL SITUATION IN RICE TO HAVE ARRIVED AT THE OPTIMUM PLANT
HEIGHT
CM FOR MAXIMUM 9P
BELOW WHICH BIOMASS DECREASES FASTER
THAN HARVEST INDEX INCREASES &IGURE
3%,%#4)/. 42!)43 &/2 )-02/6).'