Drought tolerance improvement in crop plants: An integrated view from breeding to genomics

Publisher Summary The subject of salt tolerance has received a tremendous amount of attention during the past two decades. The concept of developing salt-tolerant plants, even to the degree that they can be grown with seawater, captures the imagination of both the scientific and the public sector. There is a wide range of salt tolerance in vascular plants. Several genera include species that are extreme halophytes and are ecologically competitive only under saline conditions. Crop species primarily are glycophytes, salt-sensitive, and grow well only under cultivated conditions. Domesticated species are provided optimum conditions to obtain high yields. There is currently a need to develop new direction and cohesive impetus in the area of salt-tolerant crops development. Realistic short- and long-range goals need to be established and continually pursued. The information that is known is extensive but fragmented. This chapter outlines some of the issues and strategies concerning selection and breeding for plant salt tolerance, highlights crucial advances in knowledge in recent years, and identifies some of the fundamental gaps in our understanding.

Adaptation of Plants to Salinity

Common bean (Phaseolus vulgaris L.) is grown in regions where water deficits during reproductive development significantly reduce yield. The purpose of this study was to evaluate the association of specific phenological and physiological traits with drought resistance in common bean. Five genotypes were grown under and near a rain shelter in 1988, and an additional 16 progeny lines were included in 1990. Drought stress determined by the drought intensity index was severe (0.78) in 1988 and more moderate (0.63) in 1990. Water stress reduced the expression of most traits with the exception of days to flower and leaf moisture retention capacity. Seed yield among genotypes was reduced from 22 to 71% due to drought. Yield under stress was correlated with yield under nonstress in 1990 and negatively correlated with the drought susceptibility index in 1988. Yield components which exhibited the largest differential genotypic responses to stress were pod and seed number, whereas seed size was more stable. Genotypic variation was detected in all the partitioning indexes, chiefly harvest index and relative sink strength, and the heritability estimates for these traits were high. The limited genetic variability observed among water relations traits and their role in water conservation would restrict their potential use in the selection for drought resistance in common bean. The differential correlations between phenological, biomass and partitioning traits and the indexes for yield and drought susceptibility would suggest that the most effective approach in breeding for drought resistance in common bean would be based first on selection for high geometric yield followed by selection among the high-yielding individuals for low to moderate levels of the drought susceptibility index.

Traits related to drought resistance in common bean

Publisher Summary This chapter describes the various aspects of crop water deficits. The water status of a crop plant is usually defined in terms of its water content, water potential, or the components of water potential. The simplicity of measuring water content led to its early adoption, but the diurnal and seasonal changes in dry weight make comparisons of water content at different times of day or during the season unsatisfactory. Water deficits develop inevitably as a consequence of water loss from the leaf as the stomata open to allow the uptake of carbon dioxide from the atmosphere for photosynthesis. There have been developments in both direct and indirect methods of measuring water deficits that have resulted in considerable progress in the field. The use of infrared thermometry for the measurement of crop water deficits and the use of in situ psychrometers for the measurement of water potential respectively is elaborated. It is found that the identification of the root as the site of sensing soil water deficits does not eliminate the role of turgor pressure as the transducer of water deficits, but moves the emphasis from leaf to the root. It is observed that at the whole crop level, the water use efficiency will depend not only on the transpiration efficiency of the leaves, but also on the water loss from the soil and the optimization of yield per unit of water used.

Crop Water Deficits: A Decade of Progress

In most maize-growing areas yield reductions due to drought have been observed. Drought at flowering time is, in some cases, the most damaging. In the experiment reported here, trials with F3 families, derived from a segregating F2 population, were conducted in the field under well-watered conditions (WW) and two other water-stress regimes affecting flowering (intermediate stress, IS, and severe stress, SS). Several yield components were measured on equal numbers of plants per family: grain yield (GY), ear number (ENO), kernel number (KNO), and 100-kernel weight (HKWT). Correlation analysis of these traits showed that they were not independent of each other. Drought resulted in a 60% decrease of GY under SS conditions. By comparing yield under WW and SS conditions, the families that performed best under WW conditions were found to be proportionately more affected by stress, and the yield reductions due to SS conditions were inversely proportional to the performance under drought. Moreover, no positive correlation was observed between a drought-tolerance index (DTI) and yield under WW conditions. The correlation between GY under WW and SS conditions was 0.31. Therefore, in this experiment, selection for yield improvement under WW conditions only, would not be very effective for yield improvement under drought. Quantitative trait loci (QTLs) were identified for GY, ENO and KNO using composite interval mapping (CIM). No major QTLs, expressing more then 13% of the phenotypic variance, were detected for any of these traits, and there were inconsistencies in their genomic positions across water regimes. The use of CIM allowed the evaluation of QTL-by-environment interactions (Q×E) and could thus identify “stable” QTLs CIMMYT, Apartado Postal 6-641, 06600 Mexico D.F., Mexico across drought environments. Two such QTLs for GY, on chromosomes 1 and 10, coincided with two stable QTLs for KNO. Moreover, four genomic regions were identified for the expression of both GY and the anthesis-silking interval (ASI). In three of these, the allelic contributions were for short ASI and GY increase, while for that on chromosome 10 the allelic contribution for short ASI corresponded to a yield reduction. From these results, we hypothesize that to improve yield under drought, marker-assisted selection (MAS) using only the QTLs involved in the expression of yield components appears not to be the best strategy, and neither does MAS using only QTLs involved in the expression of ASI. We would therefore favour a MAS strategy that takes into account a combination of the “best QTLs” for different traits. These QTLs should be stable across target environments, represent the largest percentage possible of the phenotypic variance, and, though not involved directly in the expression of yield, should be involved in the expression of traits significantly correlated with yield, such as ASI.

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Identification of quantitative trait loci under drought conditions in tropical maize. 2. Yield components and marker-assisted selection strategies

With a view to understanding the basis of cultivar differences in yield under drought, a wide range of cereal cultivars representing durum wheats (Triticum turgidum L.), triticales (X Tritosecale Wittmack), barleys (Hordeum vulgare), and especially tall and dwarf bread wheats (T. aestivum L.) were studied in field experiments in north-western Mexico over three seasons. Drought was created in this rain-free environment by permanently terminating irrigation at various stages before anthesis. Control treatments were well watered throughout the growing period. Detailed measurements of plant water status, leaf area and dry matter production, anthesis date, yield components and grain yield were made. This paper presents primarily the grain yield data. Drought levels were such that the mean yield of all cultivars under drought ranged from 37 to 86% of control yield, corresponding to irrigation cut-offs varying from 69 days before mean anthesis date to only 10 days before. In each experiment the grain yield under drought showed highly significant cultivar differences, which appeared consistent between years. Yields were adjusted for drought escape by using a correction factor which ranged from 2.9 to 8.5 g/m2 per day advance in flowering, being greater in experiments with less severe drought. The demonstration of linear relationships between cultivar yield and drought intensity, as indicated by the mean yield of some or all cultivars, prompted the consideration of cultivar yield under drought as the function of yield potential (Yp, yield without drought), drought susceptibility index (S), and intensity of drought. The cultivar groups showing lowest S values (most droughtresistant) were tall bread wheats and barleys; dwarf bread wheats were intermediate, and durum wheats and triticales were the most susceptible. However, because dwarf wheats have a higher yield potential (Yp) than tall bread wheats, it is suggested that, as a group, tall bread wheats would outyield dwarf wheats only under very severe drought. Also there was considerable within-group variability of S and Yp. Cultivar S values were consistent across experiments. Yield responses of tall and dwarf bread wheat groups obtained in these experiments agreed with those seen in extensive international trials under dryland conditions.

R.A. Fischer

Papers

Drought resistance in spring wheat cultivars, 1. Grain yield responses.