Abstract
Drought is a major problem for rice grown under rainfed lowland and upland conditions, but progress in breeding to improve drought resistance has been slow. This paper describes patterns of water-stress development in rice fields, reviews genetic variation in physio-morphological traits for drought resistance in rice, and suggests how knowledge of stress physiology can contribute to plant breeding programmes that aim to increase yield in water-limiting environments. To provide a basis for integrating physiological research with plant-breeding objectives we define drought resistance in terms of relative yield of genotypes. Therefore, a drought-resistant genotype will be one which has a higher grain yield than others when all genotypes are exposed to the same level of water stress. A major reason for the slow progress in breeding for drought resistance in rice is the complexity of the drought environment, which often results in the lack of clear identification of the target environment(s). There is a need to identify the relative importance of the three common drought types; early-season drought which often causes delay in transplanting, mild intermittent stress which can have a severe cumulative effect, and late stress which affects particularly late-maturing genotypes. In addition, in rainfed lowland rice, flooded and non-flooded soil conditions may alternate during the growing season, and affect nutrient availability or cause toxicity. Several drought-resistance mechanisms, and putative traits which contribute to them, have been identified for rice; important among these being drought escape via appropriate phenology, root characteristics, specific dehydration avoidance and tolerance mechanisms, and drought recovery. Some of these mechanisms/traits have been shown to confer drought resistance and others show potential to do so in rice. The most important is the appropriate phenology which matches crop growth and development with the water environment. A deep root system, with high root length density at depth is useful in extracting water thoroughly in upland conditions, but does not appear to offer much scope for improving drought resistance in rainfed lowland rice where the development of a hard pan may prevent deep root penetration. Under water-limiting environments, genotypes which maintain the highest leaf water potential generally grow best, but it is not known if genotypic variation in leaf water potential is solely caused by root factors. Osmotic adjustment is promising, because it can potentially counteract the effects of a rapid decline in leaf water potential and there is large genetic variation for this trait. There is genotypic variation in expression of green leaf retention which appears to be a useful character for prolonged droughts, but it is affected by plant size which complicates its use as a selection criterion for drought resistance. There is a general lack of drought related research for rice in rainfed lowland conditions. This needs to be rectified, particularly considering their importance relative to upland conditions in Asian countries. We suggest that focussing physiological-genetic research efforts onto clearly defined, major target environments should provide a basis for increasing the relevance of stress physiology and the efficiency of breeding programmes for development of drought-resistant genotypes.
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