Understanding the effect of heat stress during seed filling on nutritional composition and seed yield in chickpea (Cicer arietinum L.).

Increasing temperature affects all food crops, thereby reducing their yield potential. Chickpea is a cool-season food legume vital for its nutritive value, but it is sensitive to high temperatures (> 32/20 °C maximum/minimum) during its reproductive and seed-filling stages. This study evaluated the effects of heat stress on yield and qualitative traits of chickpea seeds in a controlled environment. Chickpea genotypes differing in heat sensitivity [two heat-tolerant (HT) and two heat-sensitive (HS)] were raised in pots, initially in an outdoor environment (average 23.5/9.9 °C maximum/minimum), until the beginning of pod set (107-110 days after sowing). At this stage, the plants were moved to a controlled environment in the growth chamber to impose heat stress (32/20 °C) at the seed-filling stage, while maintaining a set of control plants at 25/15 °C. The leaves of heat-stressed plants of the HT and HS genotypes showed considerable membrane damage, altered stomatal conductance, and reduced leaf water content, chlorophyll content, chlorophyll fluorescence, and photosynthetic ability (RuBisCo, sucrose phosphate synthase, and sucrose activities) relative to their corresponding controls. Seed filling duration and seed rate drastically decreased in heat-stressed plants of the HT and HS genotypes, severely reducing seed weight plant-1 and single seed weight, especially in the HS genotypes. Yield-related traits, such as pod number, seed number, and harvest index, noticeably decreased in heat-stressed plants and more so in the HS genotypes. Seed components, such as starch, proteins, fats, minerals (Ca, P, and Fe), and storage proteins (albumin, globulins, glutelin, and prolamins), drastically declined, resulting in poor-quality seeds, particularly in the HS genotypes. These findings revealed that heat stress significantly reduced leaf sucrose production, affecting the accumulation of various seed constituents, and leading to poor nutritional quality. The HT genotypes were less affected than the HS genotypes because of the greater stability of their leaf water status and photosynthetic ability, contributing to better yield and seed quality traits in a heat-stressed environment.

Individual and combined effects of transient drought and heat stress on carbon assimilation and seed filling in chickpea.

High temperatures and decreased rainfall are detrimental to yield in chickpea (Cicer arietinum L.), particularly during grain filling. This study aimed to (i) assess the individual and combined effects of drought and heat stress on biochemical seed-filling processes, (ii) determine genotypic differences in heat and drought tolerance, and (iii) determine any cross-tolerance. Plants were grown outdoors in the normal growing season when temperatures during seed filling were <32-20°C or were planted late (temperatures >32-20°C; heat stress). Half of the pots were kept adequately watered throughout, but water was withheld from the others from the initiation of seed filling until the relative leaf water content reached 50% of the irrigated plants (drought stress); all plants were rewatered thereafter until seed maturit. Water was withheld for 13 days (normal sowing) and 7 days (late sowing), so soil moisture decreased by 54-57%. Tests on leaves and seeds were performed after the stress. Individual and combined stress damaged membranes, and decreased cellular oxidising ability, stomatal conductance, PSII function and leaf chlorophyll content; damage was greater under combined stress. Leaf Rubisco activity increased with heat stress, decreased with drought stress and decreased severely with combined stress. Sucrose and starch concentrations decreased in all seeds through reductions in biosynthetic enzymes; reductions were greater under combined stress. These effects were more severe in heat- and drought-sensitive genotypes compared with drought-tolerant genotypes. Drought stress had a greater effect than heat stress on yield and the biochemical seed-filling mechanisms. Drought- and heat-tolerant genotypes showed partial cross-tolerance.

Heat-stress-induced reproductive failures in chickpea (Cicer arietinum) are associated with impaired sucrose metabolism in leaves and anthers.

Chickpea (Cicer arietinum L.), in its reproductive stage, is sensitive to heat stress (32/20°C or higher as day/night temperatures) with consequent substantial loss of potential yields at high temperatures. The physiological mechanisms associated with reproductive failures have not been established: they constitute the basis of this study. Here, we initially screened a large core-collection of chickpea against heat stress and identified two heat-tolerant (ICC15614, ICCV. 92944) and two heat-sensitive (ICC10685, ICC5912) genotypes. These four genotypes were sown during the normal time of sowing (November-March) and also late (February-April) to expose them to heat stress during reproductive stage (>32/20°C). The genotypes were assessed for damage by heat stress to the leaves and reproductive organs using various indicators of stress injury and reproductive function. In the heat-stressed plants, phenology accelerated as days to flowering and podding, and biomass decreased significantly. The significant reduction in pod set (%) was associated with reduced pollen viability, pollen load, pollen germination (in vivo and in vitro) and stigma receptivity in all four genotypes. Heat stress inhibited pollen function more in the sensitive genotypes than in the tolerant ones, and consequently showed significantly less pod set. Heat stress significantly reduced stomatal conductance, leaf water content, chlorophyll, membrane integrity and photochemical efficiency with a larger effect on heat-sensitive genotypes. Rubisco (carbon-fixing enzyme) along with sucrose phosphate synthase (SPS) and sucrose synthase (SS) (sucrose-synthesising enzymes) decreased significantly in leaves due to heat stress leading to reduced sucrose content. Invertase, a sucrose-cleaving enzyme, was also inhibited along with SPS and SS. The inhibition of these enzymes was significantly greater in the heat-sensitive genotypes. Concurrently, the anthers of these genotypes had significantly less SPS and SS activity and thus, sucrose content. As a result, pollen had considerably lower sucrose levels, resulting in reduced pollen function, impaired fertilisation and poor pod set in heat-sensitive genotypes.