Drought (water deficits) and heat (high temperatures) stress are the prime abiotic constraints, under the current and climate change scenario in future. Any further increase in the occurrence, and extremity of these stresses, either individually or in combination, would severely reduce the crop productivity and food security, globally. Although, they obstruct productivity at all crop growth stages, the extent of damage at reproductive phase of crop growth, mainly the seed filling phase, is critical and causes considerable yield losses. Drought and heat stress substantially affect the seed yields by reducing seed size and number, eventually affecting the commercial trait ‘100 seed weight’ and seed quality. Seed filling is influenced by various metabolic processes occurring in the leaves, especially production and translocation of photoassimilates, importing precursors for biosynthesis of seed reserves, minerals and other functional constituents. These processes are highly sensitive to drought and heat, due to involvement of array of diverse enzymes and transporters, located in the leaves and seeds. We highlight here the findings in various food crops showing how their seed composition is drastically impacted at various cellular levels due to drought and heat stresses, applied separately, or in combination. The combined stresses are extremely detrimental for seed yield and its quality, and thus need more attention. Understanding the precise target sites regulating seed filling events in leaves and seeds, and how they are affected by abiotic stresses, is imperative to enhance the seed quality. It is vital to know the physiological, biochemical and genetic mechanisms, which govern the various seed filling events under stress environments, to devise strategies to improve stress tolerance. Converging modern advances in physiology, biochemistry and biotechnology, especially the “omics” technologies might provide a strong impetus to research on this aspect. Such application, along with effective agronomic management system would pave the way in developing crop genotypes/varieties with improved productivity under drought and/or heat stresses.

Rising temperatures and drought stress limit the growth and production potential of lentil (Lens culinaris Medikus), particularly during reproductive growth and seed filling. The present study aimed to (i) investigate the individual and combined effects of heat and drought stress during seed filling, (ii) determine the response of lentil genotypes with contrasting heat and drought sensitivity, and (iii) assess any cross tolerance in contrasting genotypes. For this purpose, eight lentil genotypes (two drought-tolerant, two drought-sensitive, two heat-tolerant, two heat-sensitive) were either sown at the normal time (second week of November 2014), when the temperatures at the time of seed filling were below 30/20°C (day/night), or sown late (second week of February 2015) to impose heat stress (temperatures > 30/20°C (day/night) during reproducive growth and seed filling. Half of the pots in each sowing environment were fully watered throughout (100% field capacity) while the others had water withheld (50% of field capacity) from the start of seed filling to maturity. Both heat and drought, individually or in combination, damaged cell membranes, photosynthetic traits and water relations; the effects were more severe with the combined stress. RuBisCo and stomatal conductance increased with heat stress but decreased with drought and the combined stress. Leaf and seed sucrose decreased with each stress in conjunction with its biosynthetic enzyme, while its (sucrose) hydrolysis increased under heat and drought stress, but was inhibited due to combination of stresses. Starch increased under heat stress in leaves but decreased in seeds, but drastically declined in seeds under drought alone or in combination with heat stress. At the same time, starch hydrolysis in leaves and seeds increased resulting in an accumulation of reducing sugars. Heat stress inhibited yield traits (seed number and seed weight per plant) more than drought stress, while drought stress reduced individual seed weights more than heat stress. The combined stress severely inhibited yield traits with less effect on the drought- and heat-tolerant genotypes. Drought stress inhibited the biochemical processes of seed filling more than heat stress, and the combined stress had a highly detrimental effect. A partial cross tolerance was noticed in drought and heat-tolerant lentil genotypes against the two stresses.

Ambient temperatures are predicted to rise in the future owing to several reasons associated with global climate changes. These temperature increases can result in heat stress- a severe threat to crop production in most countries. Legumes are well-known for their impact on agricultural sustainability as well as their nutritional and health benefits. Heat stress imposes challenges for legume crops and has deleterious effects on the morphology, physiology, and reproductive growth of plants. High-temperature stress at the time of the reproductive stage is becoming a severe limitation for production of grain legumes as their cultivation expands to warmer environments and temperature variability increases due to climate change. The reproductive period is vital in the life cycle of all plants and is susceptible to high-temperature stress as various metabolic processes are adversely impacted during this phase, which reduces crop yield. Food legumes exposed to high-temperature stress during reproduction show flower abortion, pollen and ovule infertility, impaired fertilization, and reduced seed filling, leading to smaller seeds and poor yields. Through various breeding techniques, heat tolerance in major legumes can be enhanced to improve performance in the field. Omics approaches unravel different mechanisms underlying thermotolerance, which is imperative to understand the processes of molecular responses toward high-temperature stress.

Rising temperatures are proving detrimental for various agricultural crops. Cool-season legumes such as lentil (Lens culunaris Medik.) are sensitive to even small increases in temperature during the reproductive stage, hence the need to explore the available germplasm for heat tolerance as well as its underlying mechanisms. In the present study, a set of 38 core lentil accessions were screened for heat stress tolerance by sowing 2 months later (first week of January; max/min temperature >32/20°C during the reproductive stage) than the recommended date of sowing (first week of November; max/min temperature <32/20°C during the reproductive stage). Screening revealed some promising heat-tolerant genotypes (IG2507, IG3263, IG3297, IG3312, IG3327, IG3546, IG3330, IG3745, IG4258, and FLIP2009) which can be used in a breeding program. Five heat-tolerant (HT) genotypes (IG2507, IG3263, IG3745, IG4258, and FLIP2009) and five heat-sensitive (HS) genotypes (IG2821, IG2849, IG4242, IG3973, IG3964) were selected from the screened germplasm and subjected to further analysis by growing them the following year under similar conditions to probe the mechanisms associated with heat tolerance. Comparative studies on reproductive function revealed significantly higher pollen germination, pollen viability, stigmatic function, ovular viability, pollen tube growth through the style, and pod set in HT genotypes under heat stress. Nodulation was remarkably higher (1.8–22-fold) in HT genotypes. Moreover, HT genotypes produced more sucrose in their leaves (65–73%) and anthers (35–78%) that HS genotypes, which was associated with superior reproductive function and nodulation. Exogenous supplementation of sucrose to in vitro-grown pollen grains, collected from heat-stressed plants, enhanced their germination ability. Assessment of the leaves of HT genotypes suggested significantly less damage to membranes (1.3–1.4-fold), photosynthetic function (1.14–1.17-fold) and cellular oxidizing ability (1.1–1.5-fold) than HS genotypes, which was linked to higher relative leaf water content (RLWC) and stomatal conductance (gS). Consequently, HT genotypes had less oxidative damage (measured as malondialdehyde and hydrogen peroxide concentration), coupled with a higher expression of antioxidants, especially those of the ascorbate–glutathione pathway. Controlled environment studies on contrasting genotypes further supported the impact of heat stress and differentiated the response of HT and HS genotypes to varying temperatures. Our studies indicated that temperatures >35/25°C were highly detrimental for growth and yield in lentil. While HT genotypes tolerated temperatures up to 40/30°C by producing fewer pods, the HS genotypes failed to do so even at 38/28°C. The findings attributed heat tolerance to superior pollen function and higher expression of leaf antioxidants.

Terminal droughts, along with high temperatures, are becoming more frequent to strongly influence the seed development in cool-season pulses like lentil. In the present study, the lentil plants growing outdoors under natural environment were subjected to following treatments at the time of seed filling till maturity: (a) 28/23 °C day/night temperature as controls; (b) drought stressed, plants maintained at 50% field capacity, under the same growth conditions as in a; (c) heat stressed, 33/28 °C day/night temperature, under the same growth conditions as in a; and (d) drought + heat stressed, plants at 50% field capacity, 33/28 °C day/night temperature, under the same growth conditions as in (a). Both heat and drought resulted in marked reduction in the rate and duration of seed filling to decrease the final seed size; drought resulted in more damage than heat stress; combined stresses accentuated the damage to seed starch, storage proteins and their fractions, minerals, and several amino acids. Comparison of a drought-tolerant and a drought-sensitive genotype indicated the former type showed significantly less damage to various components of seeds, under drought as well as heat stress suggesting a cross tolerance, which was linked to its (drought tolerant) better capacity to retain more water in leaves and hence more photo-assimilation ability, compared with drought-sensitive genotype.