Heat-tolerant Germplasm Research Articles

Durum wheat heat tolerance loci defined via a north-south gradient.

The global production of durum wheat (Triticum durum Desf.) is hindered by a constant rise in the frequency of severe heat stress events. To identify heat-tolerant germplasm, three different germplasm panels ("discovery," "investigation," and "validation") were studied under a range of heat-stressed conditions. Grain yield (GY) and its components were recorded at each site and a heat stress susceptibility index was calculated, confirming that each 1°C temperature rise corresponds to a GY reduction in durum wheat of 4.6%-6.3%. A total of 2552 polymorphic single nucleotide polymorphisms (SNPs) defined the diversity of the first panel, while 5642 SNPs were polymorphic in the "investigation panel." The use of genome-wide association studies revealed that 36 quantitative trait loci were associated with the target traits in the discovery panel, of which five were confirmed in a "subset" tested imposing heat stress by plastic tunnels, and in the investigation panel. A study of allelic combinations confirmed that Q.icd.Heat.003-1A, Q.icd.Heat.007-1B, and Q.icd.Heat.016-3B are additive in nature and the positive alleles at all three loci resulted in a 16% higher GY under heat stress. The underlying SNPs were converted into kompetitive allele specific PCR markers and tested on the validation panel, confirming that each explained up to 9% of the phenotypic variation for GY under heat stress. These markers can now be used for breeding to improve resilience to climate change and increase productivity in heat-stressed areas.

Insights into morphological and physio-biochemical adaptive responses in mungbean (Vigna radiata L.) under heat stress.

Mungbean (Vigna radiata L. Wilczek) is an important food legume crop which contributes significantly to nutritional and food security of South and Southeast Asia. The crop thrives in hot and humid weather conditions, with an optimal temperature range of 28°-35°C, and is mainly cultivated under rainfed environments. However, the rising global temperature has posed a serious threat to mungbean cultivation. Optimal temperature is a vital factor in cellular processes, and every crop species has evolved with its specific temperature tolerance ability. Moreover, variation within a crop species is inevitable, given the diverse environmental conditions under which it has evolved. For instance, various mungbean germplasm can grow and produce seeds in extreme ambient temperatures as low as 20°C or as high as 45°C. This range of variation in mungbean germplasm for heat tolerance plays a crucial role in developing heat tolerant and high yielding mungbean cultivars. However, heat tolerance is a complex mechanism which is extensively discussed in this manuscript; and at the same time individual genotypes have evolved with various ways of heat stress tolerance. Therefore, to enhance understanding towards such variability in mungbean germplasm, we studied morphological, anatomical, physiological, and biochemical traits which are responsive to heat stress in plants with more relevance to mungbean. Understanding heat stress tolerance attributing traits will help in identification of corresponding regulatory networks and associated genes, which will further help in devising suitable strategies to enhance heat tolerance in mungbean. The major pathways responsible for heat stress tolerance in plants are also discussed.

Identification of new sources of heat tolerance in cultivated and wild tomatoes

Heat stress adversely affects tomato production worldwide. Breeding thermotolerant varieties is critical to adapt to increased heat stress but tomato has a narrow genetic base for heat tolerance. Identification of new sources of heat-tolerant germplasm is important to understand physiological and molecular mechanisms of adaptation to heat stress and to broaden the genetic base of heat tolerance in tomato. Here, we screened a germplasm collection comprised of 42 tomato entries using 19 quantitative (physiological, reproductive, and morphological) and five qualitative traits. The plants were grown under long term mild heat stress with temperatures and relative humidities ranging from 23.3 to 37 °C, and 47 to 100%, respectively. The analysis of variance revealed significant differences among the entries for all the quantitative traits. Biologically significant associations were recorded for many traits. Proportion of viable pollen was not associated with fruit set percentage. The clustering analysis groups the germplasm into five clusters with a clear distinction between Solanum lycopersicum and Solanum pimpinellifolium entries. We also observed differences between the entries based on their origins. Known thermo-tolerant entries were found in Clusters 3 (eg.: LA2662, LA3317, LA3120) and 4 (e.g.: CL5915-93D4-1-0, CLN1621L, LA2661). Lines BJ01, BJ02 from Republic of Benin, and WAC1 and ATS020 from Ghana in Cluster 3 are potential new sources of thermo-tolerance in tomato. Generation of hybrids from parents belonging to the clusters 1 to 4 identified in this study would improve heterosis for heat tolerance traits in tomato. Useful traits like high pollen viability and high inflorescence numbers per plant could be introgressed from S. pimpinellifolium entries into the cultivated tomato. Our study provides important information to improve heat tolerance in tomato.

Screening of wheat germplasm for terminal heat tolerance under hyper-arid conditions

Due to intensive cropping system, late planting of wheat is very common which causes plants to suffer due to terminal heat stress. To adapt and mitigate the adverse effect of heat on wheat, researchers want to develop heat-tolerant wheat genotypes. Therefore, a set of 11 germplasms of wheat was sown in normal period (November 25) as well as in late period (December 25) at Instructional Farm of Swami Keshwanand Rajasthan Agricultural University, Bikaner. Various physiological, gas exchange and yield attributes were measured at different crop growth stages. Results showed that days to anthesis (DTA), relative water content (RWC), membrane stability index (MSI), photosynthetic pigments, photosynthetic rate (A), stomatal conductance (gs), internal CO2 concentration (ci), water use efficiency (WUE) and yield attributes were reduced, whereas transpiration rate (E) was increased under late sown as compared to normal. Most of the physiological and yield characters showed superiority (highest/lowest) in germplasm entry no. 26 followed by 43 and 58. Heat susceptibility index (HSI) was the lowest, whereas chlorophyll stability (CSI) and heat tolerance index (HTI) were the highest recorded in entry 26 followed by 43. Heat-tolerant germplasm could be selected on the basis of high RWC, MSI, photosynthetic pigments, CSI, A, gs, ci, WUE, yield components and HTI, and low E and HSI. In the present investigation, entry no. 26 (IC279317), 43 (IC335971) and 58 (IC336816) were found the best suited germplasm under heat stress. These germplasms supposed to carry the heat-tolerant genes and may be used for further heat tolerance breeding.

Morpho-physiological diversity of barley (Hordeum vulgare L.) germplasm for heat tolerance

The aim of the present study was to examine the diversity for heat tolerance in a set of 105 accessions using some key morphophysiological attributes. The data was standardized by calculating percent change over control, and used for phonetic analysis and construction of dendograms based on distance matrix (all primary data is given as supplementary material listed in statistical analysis section of Materials and Methods). These distance matrices were then used for the unweighted pair group method with arithmetic mean (UPGMA) cluster analysis using the sequential agglomerative hierarchical nonoverlapping (SAHN) method. The cluster analysis indicated some potential attributes that could be effectively used to screen germplasm for heat tolerance. Among the growth attributes, shoot fresh and dry weight was a much stable indicator than root biomass. The alterations in the concentration of chlorophyll (Chl) b played a significantly better role in clustering of various genotypes than those in Chl a. Since leaf osmotic potential was fairly maintained in most of the barley accessions under study, the notable change in leaf water and osmotic potentials seemed to be the fundamental reason for the clustering pattern. The alterations in stomatal conductance (gs ) and intercellular CO2 concentration (Ci ) were of foremost importance for clustering gas exchange parameters. The changes in electron transport rate and nonphotochemical quenching were of major importance for clustering based on chlorophyll fluorescence. Thus, it can be concluded that the thermo-tolerance and yield of barley when grown under high temperature stress can be enhanced by improving these attributes.

An overview of heat stress in tomato (Solanum lycopersicum L.).

Heat stress has been defined as the rise of temperature for a period of time higher than a threshold level, thereby permanently affecting the plant growth and development. Day or night temperature is considered as the major limiting factor for plant growth. Earlier studies reported that night temperature is an important factor in the heat reaction of the plants. Tomato cultivars capable of setting viable fruits under night temperatures above 21 °C are considered as heat-tolerant cultivars. The development of breeding objectives is generally summarized in four points: (a) cultivars with higher yield, (b) disease resistant varieties in the 1970s, (c) long shelf-life in 1980s, and (d) nutritive and taste quality during 1990s. Some unique varieties like the dwarf “Micro-Tom”, and the first transgenic tomato (FlavrSavr) were developed through breeding; they were distributed late in the 1980s.High temperature significantly affects seed, pollen viability and root expansion. Researchers have employed different parameters to evaluate the tolerance to heat stress, including membrane thermo stability, floral characteristics (Stigma exertion and antheridia cone splitting), flower number, and fruit yield per plant. Reports on pollen viability and fruit set/plant under heat stress by comparing the pollen growth and tube development in heat-treated and non-heat-stressed conditions are available in literature. The electrical conductivity (EC) have been used to evaluate the tolerance of some tomato cultivars in vitro under heat stress conditions as an indication of cell damage due to electrolyte leakage; they classified the cultivars into three groups: (a) heat tolerant, (b) moderately heat tolerant, and (c) heat sensitive.It is important to determine the range in genetic diversity for heat tolerance in tomatoes. Heat stress experiments under field conditions offer breeders information to identify the potentially heat tolerant germplasm.

EVALUATION OF COTTON (GOSSYPIUM SPP.) GERMPLASM FOR HEAT TOLERANCE UNDER NORMAL AND LATE PLANTING TIME

The objective of this study was to determine cotton (Gossypium ssp.) germplasm for heat tolerance under normal and late planting time. For this aiming 200 cotton genotypes and five check varieties (Gloria, SG 125, Flash, Ozbek 105 and Candia) were evaluated under two different temperature regimes and experiments were conducted according to the augmented design with four blocks. Field studies were carried out at the GAP International Agricultural Research and Training Center’s experimental area in Diyarbakır, Turkey, in 2016 cotton growing season. In the study heat susceptibility index was used for discriminate to the genotypes for heat tolerance. Genotypes were classified into four groups based on the heat susceptibility index. The results of this study indicated that five cotton genotypes (TAM 139-17 ELS, CIM-240, Haridost, MNH-990 and AzGR-11835) were in highly heat tolerant, 28 genotypes were found heat tolerant, 56 genotypes were in the moderately heat tolerant and other 120 genotypes were observed susceptible for heat tolerance. Based on the heat susceptibility index, five cotton genotypes can be used as parent for heat tolerance improvement in the cotton breeding program where high temperature is a limiting factor for seed cotton yield.

Genome-Wide Association Study Reveals Genomic Regions Associated With Ten Agronomical Traits in Wheat Under Late-Sown Conditions.

Poor understanding of the genetic and molecular basis of heat tolerance component traits is a major bottleneck in designing heat tolerant wheat cultivars. The impact of terminal heat stress is generally reported in the case of late sown wheat. In this study, our aim was to identify genomic regions for various agronomic traits under late sown conditions by using genome-wide association approach. An association mapping panel of 205 wheat accessions was evaluated under late sown conditions at three different locations in India. Genotyping of the association panel revealed 15,886 SNPs, out of which 11,911 SNPs with exact physical locations on the wheat reference genome were used in association analysis. A total of 69 QTLs (10 significantly associated and 59 suggestive) were identified for ten different traits including productive tiller number (17), grain yield (14), plant height (12), grain filling rate (6), grain filling duration (5), days to physiological maturity (4), grain number (3), thousand grain weight (3), harvest index (3), and biomass (2). Out of these associated QTLs, 17 were novel for traits, namely PTL (3), GY (2), GFR (6), HI (3) and GNM (3). Moreover, five consistent QTLs across environments were identified for GY (4) and TGW (1). Also, 11 multi-trait SNPs and three hot spot regions on Chr1Ds, Chr2BS, Chr2DS harboring many QTLs for many traits were identified. In addition, identification of heat tolerant germplasm lines based on favorable alleles HD2888, IC611071, IC611273, IC75240, IC321906, IC416188, and J31-170 would facilitate their targeted introgression into popular wheat cultivars. The significantly associated QTLs identified in the present study can be further validated to identify robust markers for utilization in marker-assisted selection (MAS) for development of heat tolerant wheat cultivars.

EVALUATION OF COTTON (GOSSYPIUM SPP.) GERMPLASM FOR HEAT TOLERANCE UNDER NORMAL AND LATE PLANTING TIME

The objective of this study was to determine cotton (Gossypium ssp.) germplasm for heat tolerance under normal and late planting time. For this aiming 200 cotton genotypes and five check varieties (Gloria, SG 125, Flash, Ozbek 105 and Candia) were evaluated under two different temperature regimes and experiments were conducted according to the augmented design with four blocks. Field studies were carried out at the GAP International Agricultural Research and Training Center’s experimental area in Diyarbakır, Turkey, in 2016 cotton growing season. In the study heat susceptibility index was used for discriminate to the genotypes for heat tolerance. Genotypes were classified into four groups based on the heat susceptibility index. The results of this study indicated that five cotton genotypes (TAM 139-17 ELS, CIM-240, Haridost, MNH-990 and AzGR-11835) were in highly heat tolerant, 28 genotypes were found heat tolerant, 56 genotypes were in the moderately heat tolerant and other 120 genotypes were observed susceptible for heat tolerance. Based on the heat susceptibility index, five cotton genotypes can be used as parent for heat tolerance improvement in the cotton breeding program where high temperature is a limiting factor for seed cotton yield.

Variation for Heat Tolerance During Seed Germination in Diverse Carrot [Daucus carota (L.)] Germplasm

Carrot production is constrained by high levels of heat stress during the germination stage in many global regions. Few studies have been published evaluating the effect of heat stress on carrot seed germination or screening for genetic heat stress tolerance. The objectives of this study were to evaluate the response of diverse carrot germplasm to heat stress, identify heat-tolerant germplasm that may be used by plant breeders, and define the appropriate temperature for assessing heat tolerance in germinating carrot seed. To identify an appropriate screening temperature, three commercial hybrids and an open pollinated variety were evaluated at five temperatures (24, 32.5, 35, 37.5, and 40 °C). In preliminary studies, 35 °C was identified as the optimal temperature for screening heat tolerance of carrot seed. Cultivated and wild carrot plant introductions (PIs) (n = 270) from the U.S. Department of Agriculture (USDA) National Plant Germplasm System (NPGS) representing 41 countries, inbred lines from the USDA Agricultural Research Service (n = 15), and widely grown commercial hybrids (n = 8) were evaluated for heat tolerance under heat stress and nonstress conditions (35 °C and 24 °C, respectively) by calculating absolute decrease in percent germination (AD), inhibition index (II), relative heat tolerance (RHT), and heat tolerance index (HTI). All measurements of heat tolerance identified significant differences among accessions; AD ranged from −13.0% to 86.7%, II ranged from 35.7% to 100.0%, RHT ranged from 0 to 1.36, and HTI ranged from 0.0 to 1.45. The broad-sense heritability (H2) calculations ranged from 0.64 to 0.86 for different traits, indicating a moderately strong genetic contribution to the phenotypic variation. Several wild carrot accessions and inbred lines displayed low levels of heat tolerance, whereas cultivated accessions PI 643114 (United States), PI 652400 and PI 652403 (Turkey), PI 652208 (China), and PI 652403 (Russia) were most heat tolerant. This is the first evaluation of heritability for heat stress tolerance during carrot seed germination, the first measure of HTI, and the first correlation calculation between heat and salt tolerance during germination in carrot.

Molecular breeding approaches involving physiological and reproductive traits for heat tolerance in food crops

Heat stress is a significant threat that limits crop yield and fecundity all over the world. Prevalent strategies for heat adaptation that alter technical and management systems are inadequate to sustain yield. As such, the identification of heat-tolerant genotypes with improved yield potential is crucial. Raising tolerant and stable cultivars can be tedious as heat-stress responses are highly variable across different developmental stages. While molecular breeding has progressed in engineering heat-tolerant lines, the complexity of genetic networks and divergence of heat tolerance mechanisms is the main hindrance for plant breeders. Hence, insight into the physiological and reproductive traits associated with heat tolerance could assist in the development of strategies to screen germplasm for heat tolerance. Exploitation and use of landraces and wild relatives in breeding may enhance favorable genetic diversity in crop plants. A holistic approach to delineate molecular markers, where quantitative trait loci (QTLs) for different traits linked to heat tolerance involving physiological and reproductive traits are characterized in well-controlled field environments, may be an option for optimizing germplasm under heat stress. Here, we present an outline of the effects of heat stress and its associated tolerance mechanisms in food crops, along with some physiological, molecular and reproductive characteristics such as ‘stay-green,’ membrane thermostability, canopy temperature depression, metabolites, genes, QTLs, and pollen fertility. Further, we provide information on conventional and molecular breeding approaches as well as different selection strategies for heat stress tolerance.

Differential profiles of membrane proteins, fatty acids, and sterols associated with genetic variations in heat tolerance for a perennial grass species, hard fescue (Festuca Trachyphylla)

Cellular membrane composition controls membrane stability, affecting plant tolerance to abiotic stress. The objectives of this study were to examine differential changes in membrane constituents (sterol, fatty acid and membrane protein) in response to heat stress for a cool-season perennial grass species, hard fescue (Festuca trachyphylla) and identify major membrane constituents associated with the genetic variations in heat tolerance for two varieties contrasting in heat tolerance. Plants of heat-tolerant ‘Reliant IV’ and heat-sensitive ‘Predator’ were exposed to heat stress at 38/33°C (day/night) in growth chambers. Under heat stress, ‘Reliant IV’ showed better turf quality, greater membrane stability and higher photochemical efficiency compared to ‘Predator’. Leaf membranes of ‘Reliant IV’ showed greater increase of ethyl sterols (fucosterol, stigmasterol, sitosterol and avenasterol), and two fatty acids [oleic acid (18:1) and linoleic acid (18:2)] under heat stress compared to those for ‘Predator’. Several membrane proteins were less down-regulated or more up-regulated by heat stress in ‘Reliant IV’ than in ‘Predator’, including down-regulated proteins involved in electron transport of photosynthesis (ATP synthase subunits and cytochrome b6-f complex iron-sulfur subunit), signaling (lectin-domain containing receptor kinase A4.2), and protein modification (S-acyltransferase), and up-regulated proteins involved in carboxylation in photosynthesis (Rubisco activase beta form) and stress defense (disease resistance protein 1). These differentially accumulated sterols, fatty acids, and proteins in cellular membranes could contribute to the differential level of heat tolerance between the two varieties of hard fescues. These compounds could be used as biomarkers to select for heat-tolerant germplasm in hard fescues.

Identification of High-Temperature Tolerant Lentil (Lens culinaris Medik.) Genotypes through Leaf and Pollen Traits.

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.

GENETIC ARCHITECTURE OF RELATIVE CELL INJURY AND SOME YIELD RELATED PARAMETERS IN Gossypium hirsutum L.

Although cotton is sun loving crop but its production is adversely affected due to biotic and abiotic stress, heat stress contributed significantly in this loss around the globe. The present research work on heat tolerance in cotton is a step forward to understand the genetic basis of heat tolerance in upland cotton. The presence and identification of genetic variation for certain traits is one of the pre-requisite to start research work. There are several techniques for the screening of germplasm for heat tolerance. Relative cell injury was used for the identification of the most heat tolerant and susceptible cotton genotypes i.e. VH-259, VH-142 and DNH-40, VH-282 respectively. The identified genotypes were used for the development of F1, F2, BC1 and BC2 populations for genetic studies. The biometrical analysis revealed that selected traits i.e. relative cell injury, chlorophyll contents, boll shedding percentage, plant height, number of bolls per plant, number of seeds per boll, seed cotton yield were controlled by additive and non-additive type of gene action including epistatic effects. It is suggested that selection in these populations should be done in later generations for the development of new germplasm with enhanced heat tolerance.

Grain yield, adaptation and progress in breeding for early-maturing and heat-tolerant wheat lines in South Asia

Maintaining wheat productivity under the increasing temperatures in South Asia is a challenge. We focused on developing early maturing wheat lines as an adaptive mechanism in regions suffering from terminal heat stress and those areas that require wheat adapted to shorter cycles under continual high temperature stress. We evaluated the grain yield performance of early-maturing heat-tolerant germplasm developed by CIMMYT, Mexico at diverse locations in South Asia from 2009 to 2014 and estimated the breeding progress for high-yielding and early-maturing heat-tolerant germplasm in South Asia. Each year the trial comprised of 28 new entries, one CIMMYT check (Baj) and a local check variety. Locations were classified by mega environment (ME); ME1 being the temperate irrigated locations with terminal high temperature stress, and ME5 as hot, sub-tropical, irrigated locations. Grain yield (GY), days to heading (DTH) and plant height (PH) were recorded at each location. Effect of temperature on GY was observed in both ME1 and ME5. Across years, mean minimum temperatures in ME1 and mean maximum temperatures in ME5 during grain filling had significant negative association with GY. The ME1 locations were cooler that those in ME5 in the 5 years of evaluations and had a 1–2t/ha higher GY. A mean reduction of 20days for DTH and 20cm in PH was observed in ME5. Negative genetic correlations of −0.43 to −0.79 were observed between GY and DTH in South Asia during 2009–2014. Each year, we identified early-maturing germplasm with higher grain yield than the local checks. A positive trend was observed while estimating the breeding progress across five years for high-yielding early-maturing heat tolerant wheat compared to the local checks in South Asia. The results suggests the potential of the high-yielding early-maturing wheat lines developed at CIMMYT in improving wheat production and maintaining genetic gains in South Asia.

A QTL on the short arm of wheat (Triticum aestivum L.) chromosome 3B affects the stability of grain weight in plants exposed to a brief heat shock early in grain filling.

BackgroundMolecular markers and knowledge of traits associated with heat tolerance are likely to provide breeders with a more efficient means of selecting wheat varieties able to maintain grain size after heat waves during early grain filling.ResultsA population of 144 doubled haploids derived from a cross between the Australian wheat varieties Drysdale and Waagan was mapped using the wheat Illumina iSelect 9,000 feature single nucleotide polymorphism marker array and used to detect quantitative trait loci for heat tolerance of final single grain weight and related traits. Plants were subjected to a 3 d heat treatment (37 °C/27 °C day/night) in a growth chamber at 10 d after anthesis and trait responses calculated by comparison to untreated control plants. A locus for single grain weight stability was detected on the short arm of chromosome 3B in both winter- and autumn-sown experiments, determining up to 2.5 mg difference in heat-induced single grain weight loss. In one of the experiments, a locus with a weaker effect on grain weight stability was detected on chromosome 6B. Among the traits measured, the rate of flag leaf chlorophyll loss over the course of the heat treatment and reduction in shoot weight due to heat were indicators of loci with significant grain weight tolerance effects, with alleles for grain weight stability also conferring stability of chlorophyll (‘stay-green’) and shoot weight. Chlorophyll loss during the treatment, requiring only two non-destructive readings to be taken, directly before and after a heat event, may prove convenient for identifying heat tolerant germplasm. These results were consistent with grain filling being limited by assimilate supply from the heat-damaged photosynthetic apparatus, or alternatively, accelerated maturation in the grains that was correlated with leaf senescence responses merely due to common genetic control of senescence responses in the two organs. There was no evidence for a role of mobilized stem reserves (water soluble carbohydrates) in determining grain weight responses.ConclusionsMolecular markers for the 3B or 6B loci, or the facile measurement of chlorophyll loss over the heat treatment, could be used to assist identification of heat tolerant genotypes for breeding.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-016-0784-6) contains supplementary material, which is available to authorized users.

Genetic Variation for Heat Tolerance in Primitive Cultivated Subspecies of Triticum turgidum L.

High temperature (heat) stress is a major constraint for the productivity of crops worldwide. Wheat (Triticum spp.) is particularly vulnerable to high temperature stress compared to other crops. The principal way to mitigate heat damage is to develop heat-tolerant varieties by identifying heat-tolerant germplasm and employing the tolerant lines in breeding programs. The objectives of this study were to investigate the genetic variation for heat tolerance in primitive cultivated subspecies of tetraploid wheat Triticum turgidum L. (AABB genome) and to identify heat-tolerant genotypes. Thirty-eight tetraploid wheat accessions belonging to five subspecies collected from 19 countries were studied for heat tolerance in day/night temperatures of 36/30°C beginning at anthesis. Several lines were classified as heat-tolerant based on small reductions in seed number, single-kernel weight, and total grain weight, as well as slow chlorophyll degradation under heat stress. T. turgidum subsp. polonicum IG110572 from Algeria, T. turgidum subsp. dicoccum IG45067 from Oman, and T. turgidum subsp. dicoccum IG45393 from Eritrea had responses similar to the reference heat-tolerant hexaploid wheat, ‘Kauz’. These heat-tolerant accessions may be useful to breeding programs for the improvement of heat tolerance in tetraploid or hexaploid wheat.

Transgenic tobacco plants overexpressing a grass PpEXP1 gene exhibit enhanced tolerance to heat stress.

Heat stress is a detrimental abiotic stress limiting the growth of many plant species and is associated with various cellular and physiological damages. Expansins are a family of proteins which are known to play roles in regulating cell wall elongation and expansion, as well as other growth and developmental processes. The in vitro roles of expansins regulating plant heat tolerance are not well understood. The objectives of this study were to isolate and clone an expansin gene in a perennial grass species (Poa pratensis) and to determine whether over-expression of expansin may improve plant heat tolerance. Tobacco (Nicotiana tabacum) was used as the model plant for gene transformation and an expansin gene PpEXP1 from Poa pratensis was cloned. Sequence analysis showed PpEXP1 belonged to α-expansins and was closely related to two expansin genes in other perennial grass species (Festuca pratensis and Agrostis stolonifera) as well as Triticum aestivum, Oryza sativa, and Brachypodium distachyon. Transgenic tobacco plants over-expressing PpEXP1 were generated through Agrobacterium-mediated transformation. Under heat stress (42°C) in growth chambers, transgenic tobacco plants over-expressing the PpEXP1 gene exhibited a less structural damage to cells, lower electrolyte leakage, lower levels of membrane lipid peroxidation, and lower content of hydrogen peroxide, as well as higher chlorophyll content, net photosynthetic rate, relative water content, activity of antioxidant enzyme, and seed germination rates, compared to the wild-type plants. These results demonstrated the positive roles of PpEXP1 in enhancing plant tolerance to heat stress and the possibility of using expansins for genetic modification of cool-season perennial grasses in the development of heat-tolerant germplasm and cultivars.

Genotype × environment interaction and AMMI analysis for heat tolerance in wheat

Terminal heat, which is referred as increase in temperature during grain filling, is one of the important stress factors for wheat production. Current estimates indicate that wheat crop grown on around 13.5mha in India is affected by heat stress. In order to meet the challenges of high temperature ahead of global warming, concerted efforts are needed to evaluate germplasm for heat tolerance and identify and develop genotypes suitable for such stressed environments. The advanced wheat genotypes developed for stress and normal environments by different research centers were evaluated across 7 locations representing varied agroclimatic zones during 2007–08 and 2008–09 to study their adaptability for heat stress and non-stress environments. The additive main effects and multiplicative interaction analysis for G × E interactions revealed differences amongst locations to phenology and grain yield. Genotype RAJ 4083 developed for cultivation under late sown conditions in peninsular zone was also found adaptable to timely sown conditions. Similarly, HD 2733 a cultivar of NEPZ timely sown conditions and PBW 574 an advanced breeding line of NWPZ late sown conditions was found adapted to Peninsular zone. The cultivar RAJ 3765 showed specific adaptability to Pantnagar in NWPZ. Genotype NW 3069 developed for NEPZ timely sown conditions have shown adaptability to number of locations; timely sown conditions at Karnal and Hisar in NWPZ and Niphad in PZ. Likewise, WH 1022 developed for NEPZ late sown conditions exhibited specific adaptability to all timely sown locations in NWPZ.

High temperature tolerance in chickpea and its implications for plant improvement

Chickpea (Cicer arietinum L.) is an important food legume and heat stress affects chickpea ontogeny over a range of environments. Generally, chickpea adapts to high temperatures through an escape mechanism. However, heat stress during reproductive development can cause significant yield loss. The most important effects on the reproductive phase that affect pod set, seed set and yield are: (1) flowering time, (2) asynchrony of male and female floral organ development, and (3) impairment of male and female floral organs. While this review emphasises the importance of high temperatures &gt;30°C, the temperature range of 32–35°C during flowering also produces distinct effects on grain yield. Recent field screening at ICRISAT have identified several heat-tolerant germplasm, which can be used in breeding programs for improving heat tolerance in chickpea. Research on the impact of heat stress in chickpea is not extensive. This review describes the status of chickpea production, the effects of high temperature on chickpea, and the opportunities for genetic improvement of chickpea tolerance to high temperatures.