Heat stress in grain legumes during reproductive and grain-filling phases

Thermal stress during reproductive development and grain-filling phases is a serious threat to the quality and productivity of grain legumes. The optimum temperature range for grain legume crops is 10-36°C, above which severe losses in grain yield can occur. Various climatic models have simulated that the temperature near the earth's surface will increase (by up to 4°C) by the end of this century, which will intensify the chances of heat stress in crop plants. The magnitude of damage or injury posed by a higherature stress mainly depends on the defence response of the crop and the specific growth stage of the crop at the time of exposure to the high temperature. Heat stress affects grain development in grain legumes because it disintegrates the tapetum layer, which reduces nutrient supply to microspores leading to premature anther dehiscence; hampers the synthesis and distribution of carbohydrates to grain, curtailing the grain-filling duration leading to low grain weight; induces poor pod development and fractured embryos; all of which ultimately reduce grain yield. The most prominent effects of heat stress include a substantial reduction in net photosynthetic rate, disintegration of photosynthetic apparatus and increased leaf senescence. To curb the catastrophic effect of heat stress, it is important to improve heat tolerance in grain legumes through improved breeding and genetic engineering tools and crop management strategies. In this review, we discuss the impact of heat stress on leaf senescence, photosynthetic machinery, assimilate translocation, water relations, grain quality and development processes. Furthermore, innovative breeding, genetic, molecular and management strategies are discussed to improve the tolerance against heat stress in grain legumes.