Changing paradigms in plant breeding.
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Breeding systems are important, and often neglected, aspects of the natural biology of organisms, affecting homozygosity and thus many aspects of their biology, including levels and patterns of genetic diversity and genome evolution. Among the different plant mating systems, it is useful to distinguish two types of systems: 'sex systems', hermaphroditic versus male/female and other situations; and the 'mating systems' of hermaphroditic populations, inbreeding, outcrossing or intermediate. Evolutionary changes in breeding systems occur between closely related species, and some changes occur more often than others. Understanding why such changes occur requires combined genetical and ecological approaches. I review the ideas of some of the most important theoretical models, showing how these are based on individual selection using genetic principles to ask whether alleles affecting plants' outcrossing rates or sex morphs will spread in populations. After discussing how the conclusions are affected by some of the many relevant ecological factors, I relate these theoretical ideas to empirical data from some of the many recent breeding system studies in plant populations.
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Taken together, mineral and vitamin deficiencies affect a greater number of people in the world than does protein-energy malnutrition. If farmers could be induced to grow commonly eaten food staple crops that fortify their seeds with essential vitamins and minerals, a significant, lower cost improvement in human nutrition might occur.
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The final permanent solution to micronutrient malnutrition in developing countries is a substantial improvement in dietary quality--higher consumption of pulses, fruits, vegetables, fish and animal products that the poor already desire but cannot presently afford. Meanwhile breeding staple foods that are dense in minerals and vitamins provides a low-cost, sustainable strategy for reducing levels of micronutrient malnutrition. Getting plants to do the work of fortification, referred to as "biofortification," can reach relatively remote rural populations that conventional interventions are not now reaching and can even have benefits for increased agricultural productivity. Biofortification, thus, complements conventional interventions. The symposium articles discuss several examples of ongoing research projects to develop and disseminate nutrient-dense staple food crops and issues that remain to be resolved before successful implementation can be attained.
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Drought is the main abiotic constraint on cereal yield. Analysing physiological determinants of yield responses to water may help in breeding for higher yield and stability under drought conditions. The traits to select (either for stress escape, avoidance or tolerance) and the framework where breeding for drought stress is addressed will depend on the level and timing of stress in the targeted area. If the stress is severe, breeding under stress-free conditions may be unsuccessful and traits that confer survival may become a priority. However, selecting for yield itself under stress-alleviated conditions appears to produce superior cultivars, not only for optimum environments, but also for those characterized by frequent mild and moderate stress conditions. This implies that broad avoidance/tolerance to mild-moderate stresses is given by constitutive traits also expressed under stress-free conditions. In this paper, we focus on physiological traits that contribute to improved productivity under mild-moderate drought. Increased crop performance may be achieved through improvements in water use, water-use efficiency and harvest index. The first factor is relevant when soil water remains available at maturity or when deep-rooted genotypes access water in the soil profile that is not normally available; the two latter conditions become more important when all available water is exhausted by the end of the crop cycle. Independent of the mechanism operating, a canopy able to use more water than another would have more open stomata and therefore higher canopy temperature depression, and 13C discrimination (delta13C) in plant matter. The same traits would also seem to be relevant when breeding for hot, irrigated environments. Where additional water is not available to the crop, higher water-use efficiency (WUE) appears to be an alternative strategy to improve crop performance. In this context delta13C constitutes a simple but reliable measure of WUE. However, in contrast to lines performing better because of increased access to water, lines producing greater biomass due to superior WUE will have lower delta13C values. WUE may be modified not only through a decrease in stomatal conductance, but also through an increase in photosynthetic capacity. Harvest index is strongly reduced by terminal drought (i.e. drought during grain filling). Thus, phenological traits increasing the relative amount of water used during grain filling, or adjusting the crop cycle to the seasonal pattern of rainfall may be useful. Augmenting the contribution of carbohydrate reserves accumulated during vegetative growth to grain filling may also be worthwhile in harsh environmcnts. Alternatively, extending the duration of stem elongation without changing the timing of anthesis would increase the number of grains per spike and the harvest index without changing the amount of water utilized by the crop.
Crop gene pools have adapted to and sustained the demands of agricultural systems for thousands of years. Yet, very little is known about their content, distribution, architecture, or circuitry. The presumably shallow elite gene pools often continue to yield genetic gains while the exotic pools remain mostly untapped, uncharacterized, and underutilized. The concept and content of a crop's gene pools are being changed by advancements in plant science and technology. In the first generation of plant genomics, DNA markers have refined some perceptions of genetic variation by providing a glimpse of a primary source, DNA polymorphism. The markers have provided new and more powerful ways of assessing genetic relationships, diversity, and merit by infusing genetic information for the first time in many scenarios or in a more comprehensive manner for others. As a result, crop gene pools may be supplemented through more rapid and directed methods from a greater variety of sources. Previously limited by the barriers of sexual reproduction, the native gene pools will soon be complemented by another gene pool (transgenes) and perhaps by other native exotic gene pools through comparative analyses of plants' biological repertoire. Plant genomics will be an important force of change for crop improvement. The plant science community and crop gene pools may be united and enriched as never before. Also, the genomes and gene pools, the products of evolution and crop domestication, will be reduced and subjected to the vagaries and potential divisiveness of intellectual property considerations. Let the gains begin.
An analysis based on Elston's model of mixed major locus and polygenic inheritance is extended to include populations of progeny testing such as F3, B1s and B2s families derived from F2 and backcrosses in a cross between two inbred lines. Genetic hypotheses that can be validly tested by the likelihood ratio method in the analysis of a breeding experiment include homogeneity of variances due to environment and/or polygenes with transformable scale effect by Box-Cox power function, random and independent segregation of major genes, invariance of the effects of major genes with population types and additive and dominant models for polygenes. Testing hypotheses in the order suggested here can lead to a gradual simplification of the models and increases the feasibility of the subsequent analysis, but caution must be paid to the possible bias in parameter estimation and hypotheses tests. The procedure is applied to a set of data on plant height of rice with the effects of dwarf genes in crosses among three varieties. Two recessive dwarf genes are shown to be nonallelic and unlinked. One dwarf gene is shown to reduce plant height about 36-56 cm, and another 52-61 cm. The effect of polygenes, estimated as the standard deviation among possible inbred lines derived from these crosses, is about 11.7 cm. Interactions between the dwarf genes and the polygenic background are found, especially for one of the two genes. Both the polygenic effects and the interactions are much smaller than the effects of the major dwarf genes.
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A new index of selection efficiency, i.e. the ratio of the probability of achieving the desired genetic gain in a target population to the cost expended for that gain, is introduced to define the optimum selection procedures that give the most opportunities for producing new commercial varieties. Monte Carlo simulations of mass selection showed that the index gives solutions to some important optimization problems which cannot be solved based on the traditional indices, the expected genetic gain or its ratio to the cost. The optimum cycle and intensity of selection and optimum population size for a target population could be defined by the new index, but not by the traditional indices.
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