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Abstract 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.

Rising temperatures impact different developmental stages of summer crops like mung bean, particularly during the crucial seed-filling stage. This study focused on two mung bean genotypes, categorized as heat-tolerant [HT] or heat-sensitive [HS]. These genotypes were grown in pots in an outdoor natural environment (average day/night temperature 36°C/24.3°C) until the onset of podding (40 days after sowing) and subsequently relocated to controlled-environment walk-in growth chambers for exposure to heat stress (42°C/30°C) or control conditions (35°C/25°C) until maturity. For all measured attributes, heat stress had a more pronounced effect on the HS genotype than on the HT genotype. Heat-stressed plants exhibited severe leaf damage, including membrane damage, reduced chlorophyll content, diminished chlorophyll fluorescence, and decreased leaf water content. Heat stress impeded the seed-filling rate and duration, decreasing starch, protein, fat, and mineral contents, with a notable decline in storage proteins. Heat stress disrupted the activities of several seed enzymes, inhibiting starch and sucrose accumulation and consequently decreasing individual seed weights and seed weight plant−1. This study revealed that heat stress during seed filling severely impaired mung bean seed yield and nutritional quality due to its impact on various stress-related traits in leaves and enzyme activities in seeds. Moreover, this research identified potential mechanisms related to heat tolerance in genotypes with contrasting heat sensitivity.

The rising temperatures are seriously impacting the food crops, including urdbean; hence efforts are needed to identify the sources of heat tolerance in such crops to ensure global food security. In the present study, urdbean genotypes were evaluated for heat tolerance under natural outdoor for two consecutive years (2018, 2019) and subsequently in the controlled environment of the growth chamber to identify high temperature tolerant lines. The genotypes were assessed involving few physiological traits (membrane damage, chlorophyll, photosynthetic efficiency, stomatal conductance, lipid peroxidation), reproductive traits (pollen germination % and pollen viability %) and yield related traits (total number of pods plant-1, total seeds plant-1, single seed weight and seed yield plant-1). Based upon these tested traits, PantU31, Mash114, UTTARA and IPU18-04 genotypes were identified as promising genotypes for both years under heat stress condition. Further confirming heat tolerance, all these four tolerant and four sensitive genotypes were tested under controlled environment under growth chamber condition. All these four genotypes PantU31, Mash114, UTTARA and IPU18-04 showed high chlorophyll content, photosynthetic efficiency, stomatal conductance, leaf area, pods plant-1, total seeds plant-1 and low reduction in pollen germination % and pollen viability under stress heat stress condition. Moreover, yield and yield related traits viz., pods plant-1, seeds plant-1, single seed weight and seed yield plant-1 showed very strong positive correlation with pollen germination and pollen viability except electrolyte leakage and malondialdehyde content. Thus, these genotypes could be potentially used as donors for transferring heat tolerance trait to the elite yet heat-sensitive urdbean cultivars.

Vegetables are a distinct collection of plant-based foods that vary in nutritional diversity and form an important part of the healthy diet of the human being. Besides providing basic nutrition, they have great potential for boosting human health. The balanced consumption of vegetables is highly recommended for supplementing the human body with better nutrition density, dietary fiber, minerals, vitamins, and bioactive compounds. However, the production and quality of fresh vegetables are influenced directly or indirectly by exposure to high temperatures or heat stress (HS). A decline in quality traits and harvestable yield are the most common effects of HS among vegetable crops. Heat-induced morphological damage, such as poor vegetative growth, leaf tip burning, and rib discoloration in leafy vegetables and sunburn, decreased fruit size, fruit/pod abortion, and unfilled fruit/pods in beans, are common, often rendering vegetable cultivation unprofitable. Further studies to trace down the possible physiological and biochemical effects associated with crop failure reveal that the key factors include membrane damage, photosynthetic inhibition, oxidative stress, and damage to reproductive tissues, which may be the key factors governing heat-induced crop failure. The reproductive stage of plants has extensively been studied for HS-induced abnormalities. Plant reproduction is more sensitive to HS than the vegetative stages, and affects various reproductive processes like pollen germination, pollen load, pollen tube growth, stigma receptivity, ovule fertility and, seed filling, resulting in poorer yields. Hence, sound and robust adaptation and mitigation strategies are needed to overcome the adverse impacts of HS at the morphological, physiological, and biochemical levels to ensure the productivity and quality of vegetable crops. Physiological traits such as the stay-green trait, canopy temperature depression, cell membrane thermostability, chlorophyll fluorescence, relative water content, increased reproductive fertility, fruit numbers, and fruit size are important for developing better yielding heat-tolerant varieties/cultivars. Moreover, various molecular approaches such as omics, molecular breeding, and transgenics, have been proved to be useful in enhancing/incorporating tolerance and can be potential tools for developing heat-tolerant varieties/cultivars. Further, these approaches will provide insights into the physiological and molecular mechanisms that govern thermotolerance and pave the way for engineering “designer” vegetable crops for better health and nutritional security. Besides these approaches, agronomic methods are also important for adaptation, escape and mitigation of HS protect and improve yields.

Under global climate change, high-temperature stress is becoming a major threat to crop yields, adversely affecting plant growth, and ultimately resulting in significant yield losses in various crops, including chickpea. Thus, identifying crop genotypes with increased heat stress (HS) tolerance is becoming a priority for chickpea research. Here, we assessed the response of seven physiological traits and four yield and yield-related traits in 39 chickpea genotypes grown in normal-sown and late-sown environments [to expose plants to HS (>32/20°C) at the reproductive stage] for two consecutive years (2017–2018 and 2018–2019). Significant genetic variability for the tested traits occurred under normal and HS conditions in both years. Based on the tested physiological parameters and yield-related traits, GNG2171, GNG1969, GNG1488, PantG186, CSJ515, RSG888, RSG945, RVG202, and GNG469 were identified as promising genotypes under HS. Further, ten heat-tolerant and ten heat-sensitive lines from the set of 39 genotypes were validated for their heat tolerance (32/20°C from flowering to maturity) in a controlled environment of a growth chamber. Of the ten heat-tolerant genotypes, GNG1969, GNG1488, PantG186, RSG888, CSJ315, and GNG1499 exhibited high heat tolerance evidenced by small reductions in pollen viability, pollen germination, and pod set %, high seed yield plant–1 and less damage to membranes, photosynthetic ability, leaf water status, and oxidative processes. In growth chamber, chlorophyll, photosynthetic efficiency, pollen germination, and pollen viability correlated strongly with yield traits. Thus, GNG1969, GNG1488, PantG186, RSG888, CSJ315, and GNG1499 genotypes could be used as candidate donors for transferring heat tolerance traits to high-yielding heat-sensitive varieties to develop heat-resilient chickpea cultivars.

Grain legumes are a key food source for ensuring global food security and sustaining agriculture. However, grain legume production is challenged by growing disease incidence due to global climate change. Ascochyta blight (AB) is a major disease, causing substantial yield losses in grain legumes worldwide. Harnessing the untapped reserve of global grain legume germplasm, landraces, and crop wild relatives (CWRs) could help minimize yield losses caused by AB infection in grain legumes. Several genetic determinants controlling AB resistance in various grain legumes have been identified following classical genetic and conventional breeding approaches. However, the advent of molecular markers, biparental quantitative trait loci (QTL) mapping, genome-wide association studies, genomic resources developed from various genome sequence assemblies, and whole-genome resequencing of global germplasm has revealed AB-resistant gene(s)/QTL/genomic regions/haplotypes on various linkage groups. These genomics resources allow plant breeders to embrace genomics-assisted selection for developing/transferring AB-resistant genomic regions to elite cultivars with great precision. Likewise, advances in functional genomics, especially transcriptomics and proteomics, have assisted in discovering possible candidate gene(s) and proteins and the underlying molecular mechanisms of AB resistance in various grain legumes. We discuss how emerging cutting-edge next-generation breeding tools, such as rapid generation advancement, field-based high-throughput phenotyping tools, genomic selection, and CRISPR/Cas9, could be used for fast-tracking AB-resistant grain legumes to meet the increasing demand for grain legume-based protein diets and thus ensuring global food security.

Rising global temperatures due to climate change are affecting crop performance in several regions of the world. High temperatures affect plants at various organizational levels, primarily accelerating phenology to limit biomass production and shortening reproductive phase to curtail flower and fruit numbers, thus resulting in severe yield losses. Besides, heat stress also disrupts normal growth, development, cellular metabolism, and gene expression, which alters shoot and root structures, branching patterns, leaf surface and orientation, and anatomical, structural, and functional aspects of leaves and flowers. The reproductive growth stage is crucial in plants’ life cycle, and susceptible to high temperatures, as reproductive processes are negatively impacted thus reducing crop yield. Genetic variation exists among genotypes of various crops to resist impacts of heat stress. Several screening studies have successfully phenotyped large populations of various crops to distinguish heat-tolerant and heat-sensitive genotypes using various traits, related to shoots (including leaves), flowers, fruits (pods, spikes, spikelets), and seeds (or grains), which have led to direct release of heat-tolerant cultivars in some cases (such as chickpea). In the present review, we discuss examples of contrasting genotypes for heat tolerance in different crops, involving many traits related to thermotolerance in leaves (membrane thermostability, photosynthetic efficiency, chlorophyll content, chlorophyll fluorescence, stomatal activity), flowers (pollen viability, pollen germination, fertilization, ovule viability), roots (architecture), biomolecules (antioxidants, osmolytes, phytohormones, heat-shock proteins, other stress proteins), and “omics” (phenomics, transcriptomics, genomics) approaches. The traits linked to heat tolerance can be introgressed into high yielding but heat-sensitive genotypes of crops to enhance their thermotolerance. Involving these traits will be useful for screening 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.

Chickpea is one of the most economically important food legumes, and a significant source of proteins. It is cultivated in more than 50 countries across Asia, Africa, Europe, Australia, North America, and South America. Chickpea production is limited by various abiotic stresses (cold, heat, drought, salt, etc.). Being a winter-season crop in northern south Asia and some parts of the Australia, chickpea faces low-temperature stress (0–15°C) during the reproductive stage that causes substantial loss of flowers, and thus pods, to inhibit its yield potential by 30–40%. The winter-sown chickpea in the Mediterranean, however, faces cold stress at vegetative stage. In late-sown environments, chickpea faces high-temperature stress during reproductive and pod filling stages, causing considerable yield losses. Both the low and the high temperatures reduce pollen viability, pollen germination on the stigma, and pollen tube growth resulting in poor pod set. Chickpea also experiences drought stress at various growth stages; terminal drought, along with heat stress at flowering and seed filling can reduce yields by 40–45%. In southern Australia and northern regions of south Asia, lack of chilling tolerance in cultivars delays flowering and pod set, and the crop is usually exposed to terminal drought. The incidences of temperature extremes (cold and heat) as well as inconsistent rainfall patterns are expected to increase in near future owing to climate change thereby necessitating the development of stress-tolerant and climate-resilient chickpea cultivars having region specific traits, which perform well under drought, heat, and/or low-temperature stress. Different approaches, such as genetic variability, genomic selection, molecular markers involving quantitative trait loci (QTLs), whole genome sequencing, and transcriptomics analysis have been exploited to improve chickpea production in extreme environments. Biotechnological tools have broadened our understanding of genetic basis as well as plants' responses to abiotic stresses in chickpea, and have opened opportunities to develop stress tolerant chickpea.

Chickpea is sensitive to cold stress, especially at reproductive stage, resulting in flower and pod abortion that significantly reduces seed yield. In the present study, we evaluated (a) whether cold acclimation imparts reproductive cold tolerance in chickpea; (b) how genotypes with contrasting sensitivity respond to cold acclimation; and (c) the involvement of cryoprotective solutes and antioxidants in anthers and ovules in cold acclimation. Four chickpea genotypes with contrasting cold sensitivity (cold-tolerant: ICC 17258, ICC 16349; cold-sensitive: ICC 15567, GPF 2) were grown in an outdoor environment for 40 days in November (average maximum/minimum temperature 24.9/15.9 °C) before being subjected to cold stress (13/7 °C), with or without cold acclimation in a controlled environment of walk-in-growth chambers. The 42-d cold acclimation involved 7 d exposure at each temperature beginning with 23/15 °C, 21/13 °C, 20/12 °C, 20/10 °C, 18/8 °C, 15/8 °C (12 h/12 h day/night), prior to exposing the plants to cold stress (13/7 °C, 12 h/12 h day/night; 700 μmol m−2 s−1 light intensity; 65–70% relative humidity). Cold acclimation remarkably reduced low temperature-induced leaf damage (as membrane integrity, leaf water status, stomatal conductance, photosynthetic pigments, and chlorophyll fluorescence) under cold stress in all four genotypes. It only reduced anther and ovule damage in cold-tolerant genotypes due to improved antioxidative ability, measured as enzymatic (superoxide dismutase, catalase, ascorbate peroxidase, and glutathione reductase) and non-enzymatic (ascorbate and reduced glutathione), solutes (particularly sucrose and γ-aminobutyric acid) leading to improving reproductive function and yield traits, whereas cold-sensitive genotypes were not responsive. The study concluded that cold tolerance in chickpea appears to be related to the better ability of anthers and ovules to acclimate, involving various antioxidants and cryoprotective solutes. This information will be useful in directing efforts toward increasing cold tolerance in chickpea.

Cold stress modifies anthers’ metabolic pathways to induce pollen sterility. Cold-tolerant plants, unlike the susceptible ones, produce high proportion of viable pollen. Anthers in susceptible plants, when exposed to cold stress, increase abscisic acid (ABA) metabolism and reduce ABA catabolism. Increased ABA negatively regulates expression of tapetum cell wall bound invertase and monosaccharide transport genes resulting in distorted carbohydrate pool in anther. Cold-stress also reduces endogenous levels of the bioactive gibberellins (GAs), GA4 and GA7, in susceptible anthers by repression of the GA biosynthesis genes. Here we discuss recent findings on mechanisms of cold susceptibility in anthers which determine pollen sterility. We also discuss differences in regulatory pathways between cold-stressed anthers of susceptible and tolerant plants that decide pollen sterility or viability.