Chlorophyll Stability is an Indicator of Drought Tolerance in Peanut

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.

Following the recognition of the importance of dealing with the effects of genotype-by-environment (G ×E) interaction in multi-environment testing of genotypes in plant breeding programs, there has been substantial development in the area of analytical methodology to quantify and describe these interactions. Three major areas where there have been developments are the analysis of variance, indirect selection, and pattern analysis methodologies. This has resulted in a wide range of analytical methods each with their own advocates. There is little doubt that the development of these methodologies has greatly contributed to an enhanced understanding of the magnitude and form ofG ×E interactions and our ability to quantify their presence in a multi-environment experiment. However, our understanding of the environmental and physiological bases of the nature ofG ×E interactions in plant breeding has not improved commensurably with the availability of these methodologies. This may in part be due to concentration on the statistical aspects of the analytical methodologies rather than on the complementary resolution of the biological basis of the differences in genotypic adaptation observed in plant breeding experiments. There are clear relationships between many of the analytical methodologies used for studying genotypic variation andG ×E interaction in plant breeding experiments. However, from the numerous discussions on the relative merits of alternative ways of analysingG ×E interactions which can be found in the literature, these relationships do not appear to be widely appreciated. This paper outlines the relevant theoretical relationships between the analysis of variance, indirect selection and pattern analysis methodologies, and their practical implications for the plant breeder interested in assessing the effects ofG ×E interaction on the response to selection. The variance components estimated from the combined analysis of variance can be used to judge the relative magnitude of genotypic andG ×E interaction variance. Where concern is on the effect of lack of correlation among environments, theG ×E interaction component can be partitioned into a component due to heterogeneity of genotypic variance among environments and another due to the lack of correlation among environments. In addition, the pooled genetic correlation among all environments can be estimated as the intraclass correlation from the variance components of the combined analysis of variance. WhereG ×E interaction accounts for a large proportion of the variation among genotypes, the individual genetic correlations between environments could be investigated rather than the pooled genetic correlation. Indirect selection theory can be applied to the case where the same character is measured on the same genotypes in different environments. Where there are no correlations of error effects among environments, the phenotypic correlation between environments may be used to investigate indirect response to selection. Pattern analysis (classification and ordination) methods based on standardised data can be used to summarise the relationships among environments in terms of the scope to exploit indirect selection. With the availability of this range of analytical methodology, it is now possible to investigate the results of more comprehensive experiments which attempt to understand the nature of differences in genotypic adaptation. Hence a greater focus of interest on understanding the causes of the interaction can be achieved.