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.

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.

The mechanisms affecting the heat sensitivity of chickpea are largely unknown. Heat-tolerant (ICCV07110, ICCV92944) and heat-sensitive (ICC14183, ICC5912) chickpea genotypes were sown in February in the soil-filled pots. At the time of flowering, these were subjected to varying day/night temperatures of 30/20, 35/25, 40/30 and 45/358C in the growth chambers (12 h light/12 h dark; light intensity, 250 mmol m 72 s 71 , 80% relative humidity). The pollen viability, pollen germination, tube growth, pollen load and stigma receptivity decreased with increases in temperatures to 45/358C. The heat-tolerant genotypes experienced significantly less damage to pollen and stigma function. Membrane integrity, chlorophyll content, photochemical efficiency and cellular oxidizing ability were inhibited by the increase in temperature, with greater impacts on the sensitive genotypes. Oxidative injury as lipid peroxidation and hydrogen peroxide content was significantly greater in sensitive genotypes at 40/30 and 45/358C. Enzymatic and non-enzymatic antioxidants showed increased levels at 40/308C, but decreased considerably at 45/358C. Heat-tolerant genotypes possessed greater activity of ascorbate peroxidase and glutathione reductase, along with higher levels of ascorbate and reduced glutathione at 40/30 and 45/ 358C. Biomass, pod set and yield were not affected significantly at 35/258C, but began to decrease at 40/308C and were lowest at 45/358C. The sensitive genotypes were not able to set any pods at 45/358C, whereas the tolerant genotypes produced only few fertile pods at this temperature. It was concluded that heat stress leads to loss of pollen as well as stigma function and induces oxidative stress in the leaves that cause failure of fertilization and damage to the leaves, respectively.

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