Heat Tolerance in Crops contrasting genotypes and would pave the way for characterizing the underlying molecular mechanisms, which could be valuable for engineering plants with enhanced thermotolerance. Wherever possible, we discussed breeding and biotechnological approaches for using these traits to develop heat-tolerant genotypes of various food crops.

Increasing temperatures are adversely affecting various food crops, including legumes, and this issue requires attention. The growth of two cool-season food legumes, chickpea and lentil, is inhibited by high temperatures but their relative sensitivity to heat stress and the underlying reasons have not been investigated. Moreover, the high-temperature thresholds for these two legumes have not been well-characterised. In the present study, three chickpea (ICCVO7110, ICC5912 and ICCV92944) and two lentil (LL699 and LL931) genotypes, having nearly similar phenology with respect to flowering, were grown at 30/20°C (day/night; control) until the onset of flowering and subsequently exposed to varying high temperatures (35/25, 38/28, 40/30 and 42/32°C; day/night) in a controlled environment (growth chamber; 12 hr/12 hr; light intensity 750 µmol m −2 s −1 ; RH-70%) at 108 days after sowing for both the species. Phenology (podding, maturity) was accelerated in both the species; the days to podding declined more in lentil at 35/25 (2.8 days) and 38/28°C (11.3 days) than in chickpea (1.7 and 7.1 days, respectively). Heat stress decreased flowering-podding and podding-maturity intervals considerably in both the species. At higher temperatures, no podding was observed in lentil, while chickpea showed reduction of 14.9 and 16.1 days at 40/30 and 42/32°C, respectively. Maturity was accelerated on 15.3 and 12.5 days at 38/28°C, 33.6 and 34 days at 40/30°C and 45.6 and 47 days at 42/32°C, in chickpea and lentil, respectively. Consequently, biomass decreased considerably at 38/28°C in both the species to limit the yield-related traits. Lentil was significantly more sensitive to heat stress, with the damage-assessed as reduction in biomass, reproductive function-related traits (pollen viability, germination, pollen tube growth and stigma receptivity), leaf traits such as membrane injury, leaf water status, photochemical efficiency, chlorophyll concentration, carbon fixation and assimilation, and oxidative stress, appearing even at 35/25°C, compared with 38/28°C, in chickpea. The expression of enzymatic antioxidants such as superoxide dismutase, catalase, ascorbate peroxidase, glutathione reductase and non-enzymatic | 735 BHANDARI et Al. 1 | INTRODUC TI ON Rising temperatures, both globally and locally, are resulting in heat stress for various summer-and cool-season food crops (Sehgal et al., 2018). Heat stress for most subtropical and tropical crops refers to situations where the temperature exceeds 32-35°C (maximum value; Bita & Gerats, 2013); however, for cool-season crops, a daily maximum temperature more than 25°C (maximum value) is considered stressful (Wahid, Gelani, Ashraf, & Foolad, 2007; Wardlaw & Wrigley, 1994). The effect of heat stress varies depending on the temperature, duration of exposure and intensity (Wahid et al., 2007). Heat stress accelerates phenology and causes a significant decline in vegetative and reproductive duration (pod and seed number, seed filling) to inhibit the potential yields, as reported in various

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Gradually increasing temperatures at global and local scales are causing heat stress for cool and summer-season food legumes, such as lentil (Lens culinaris Medik.), which is highly susceptible to heat stress, especially during its reproductive stages of development. Hence, suitable strategies are needed to develop heat tolerance in this legume. In the present study, we tested the effectiveness of heat priming (HPr; 6 h at 35 °C) the lentil seeds and a foliar treatment of γ-aminobutyric acid (GABA; 1 mM; applied twice at different times), singly or in combination (HPr+GABA), under heat stress (32/20 °C) in two heat-tolerant (HT; IG2507, IG3263) and two heat-sensitive (HS; IG2821, IG2849) genotypes to mitigate heat stress. The three treatments significantly reduced heat injury to leaves and flowers, particularly when applied in combination, including leaf damage assessed as membrane injury, cellular oxidizing ability, leaf water status, and stomatal conductance. The combined HPr+GABA treatment significantly improved the photosynthetic function, measured as photosynthetic efficiency, chlorophyll concentration, and sucrose synthesis; and significantly reduced the oxidative damage, which was associated with a marked up-regulation in the activities of enzymatic antioxidants. The combined treatment also facilitated the synthesis of osmolytes, such as proline and glycine betaine, by upregulating the expression of their biosynthesizing enzymes (pyrroline-5-carboxylate synthase; betaine aldehyde dehydrogenase) under heat stress. The HPr+GABA treatment caused a considerable enhancement in endogenous levels of GABA in leaves, more so in the two heat-sensitive genotypes. The reproductive function, measured as germination and viability of pollen grains, receptivity of stigma, and viability of ovules, was significantly improved with combined treatment, resulting in enhanced pod number (21–23% in HT and 35–38% in HS genotypes, compared to heat stress alone) and seed yield per plant (22–24% in HT and 37–40% in HS genotypes, in comparison to heat stress alone). The combined treatment (HPr+GABA) was more effective and pronounced in heat-sensitive than heat-tolerant genotypes for all the traits tested. This study offers a potential solution for tackling and protecting heat stress injury in lentil plants.

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Rising temperatures, globally and locally, would be detrimental for cool-and summer-season food legumes, such as lentil (Lens culinaris Medik.). Lentil is highly sensitive to supra-optimal temperatures ([ 30 °C), particularly during reproductive growth, resulting in flower and pod losses. Thus, suitable strategies are needed to introduce heat tolerance in this legume. Here, we evaluated the efficacy of nitric oxide (NO)-applied as foliar treatment of 1 mM sodium nitroprusside (SNP), twice (one day before final exposure to high temperature, and again five days later)-on heat-stressed (32/20 °C) lentil genotypes, differing in heat sensitivity. As a result of heat stress, endogenous NO increased significantly in heat-tolerant genotypes (46-62% in leaves and 66-68% in anthers, relative to the respective controls), while it decreased in heatsensitive (HS) genotypes (27-30% in leaves and 28-33% in anthers, relative to the respective controls). Foliar supplementation with SNP markedly increased endogenous NO in leaves and anthers of both the control and heattreated plants. Heat stress significantly accelerated phenology, damaged membranes, chlorophyll, chlorophyll fluorescence, cellular viability, and decreased leaf water status, carbon fixing and assimilating ability, less so in plants treated with SNP. Heat stress plus SNP significantly improved carbon fixation (as RuBisCo activity) and assimilation ability, (as sucrose concentration (in leaves and anthers), sucrose synthase and vacuolar acid invertase activity, reducing sugars), as well as osmolyte accumulation (proline and glycine betaine) in leaves and anthers. Moreover, SNP-treated plants had significantly less oxidative damage-measured as malondialdehyde and hydrogen peroxide concentrations-in leaves and anthers, relative to the respective control. Reproductive functionassessed as pollen grain germination and viability, stigma receptivity, and ovular viability-decreased markedly in plants exposed to heat stress alone, more so in HS genotypes, but increased significantly with SNP treatment as a consequence of improved leaf and anther function, to significantly increase the pod and seed numbers in heatstressed lentil plants, relative to heat-stress alone.