Heat Stress in Wheat during Reproductive and Grain-Filling Phases

Ambient temperatures have increased since the beginning of the century and are predicted to continue rising under climate change. Such increases in temperature can cause heat stress: a severe threat to wheat production in many countries, particularly when it occurs during reproductive and grain-filling phases. Heat stress reduces plant photosynthetic capacity through metabolic limitations and oxidative damage to chloroplasts, with concomitant reductions in dry matter accumulation and grain yield. Genotypes expressing heat shock proteins are better able to withstand heat stress as they protect proteins from heat-induced damage. Heat tolerance can be improved by selecting and developing wheat genotypes with heat resistance. Wheat pre-breeding and breeding may be based on secondary traits like membrane stability, photosynthetic rate and grain weight under heat stress. Nonetheless, improvement in grain yield under heat stress implies selecting genotypes for grain size and rate of grain filling. Integrating physiological and biotechnological tools with conventional breeding techniques will help to develop wheat varieties with better grain yield under heat stress during reproductive and grain-filling phases. This review discusses the impact of heat stress during reproductive and grain-filling stages of wheat on grain yield and suggests strategies to improve heat stress tolerance in wheat.     

Productivity of a doubled haploid winter wheat population under heat stress

Breeding of new winter wheat cultivars with good heat tolerance requires better understanding of the genetic background of heat tolerance. In the present work the effect of heat stress on the 6^th day after heading was investigated in a doubled haploid (DH) population arising from a cross between heat-sensitive (Plainsman V) and heat-tolerant (Mv Magma) cultivars. Averaged over the population, heat stress was found to result in a significant reduction in biomass, grain yield and grain number per plant, and in thousand-kernel weight (TKW) and harvest index. High temperature had the greatest effect on the grain yield, with a drop of 36.2% compared with the control. This could be attributed jointly to significant reductions in the TKW of the main ear and in the grain number of the side tillers. The relationship between the yield parameters was confirmed by the positive correlations obtained for the lines in the population. The diverse levels of heat tolerance in the different lines were confirmed by the significant differences in the reduction in the chlorophyll content (SPAD index) of the flag-leaves and in yield parameters. The changes in yield components in stress condition, however, can be still the most effective tools for heat stress evaluation.     

QTL for yield and associated traits in the Seri/Babax population grown across several environments in Mexico, in the West Asia, North Africa, and South Asia regions

Heat and drought adaptive quantitative trait loci (QTL) in a spring bread wheat population resulting from the Seri/Babax cross designed to minimize confounding agronomic traits have been identified previously in trials conducted in Mexico. The same population was grown across a wide range of environments where heat and drought stress are naturally experienced including environments in Mexico, West Asia, North Africa (WANA), and South Asia regions. A molecular genetic linkage map including 475 marker loci associated to 29 linkage groups was used for QTL analysis of yield, days to heading (DH) and to maturity (DM), grain number (GM2), thousand kernel weight (TKW), plant height (PH), canopy temperature at the vegetative and grain filling stages (CTvg and CTgf), and early ground cover. A QTL for yield on chromosome 4A was confirmed across several environments, in subsets of lines with uniform allelic expression of a major phenology QTL, but not independently from PH. With terminal stress, TKW QTL was linked or pleiotropic to DH and DM. The link between phenology and TKW suggested that early maturity would favor the post—anthesis grain growth periods resulting in increased grain size and yields under terminal stress. GM2 and TKW were partially associated with markers at different positions suggesting different genetic regulation and room for improvement of both traits. Prediction accuracy of yield was improved by 5 % when using marker scores of component traits (GM2 and DH) together with yield in multiple regression. This procedure may provide accumulation of more favorable alleles during selection.     

不同生态环境与播种期下小麦籽粒品质变异规律的研究

选用 6个不同品质类型小麦品种在 4个生态点进行分期播种试验, 系统分析了不同生态环境下小麦籽粒产量与品质的变异特征及其与主要气候生态因子间的关系。结果表明 :生态点、品种以及地点×品种互作对籽粒产量、千粒重、蛋白质、湿面筋和淀粉含量、沉降值与降落值的影响均达到显著水平 ;不同播种期处理对产量与淀粉含量的影响达到极显著水平, 而播种期×品种互作对千粒重、降落值、淀粉含量及沉降值的效应达到显著水平 ;地点×播种期×品种互作仅对产量、湿面筋、淀粉含量与沉降值有显著的影响。在 4个不同生态点中, 南京点的籽粒蛋白质与湿面筋含量最低, 但淀粉含量最高 ;徐州点的产量和千粒重最大 ;泰安点的蛋白质含量与湿面筋含量最高, 沉降值最小 ;保定点的产量、千粒重最小, 但沉降值最大。不同播种期处理下, 适播与晚播的籽粒蛋白质含量、湿面筋含量、淀粉含量、沉降值与降落值都显著高于早播, 而早播期的产量和千粒重最大。各小麦品种在不同地点与播种期下产量与品质性状的变异中以降落值的变异系数为最大, 淀粉的变异为最小。开花至成熟期的日均温与淀粉含量呈线性负相关, 与产量、蛋白质和湿面筋含量及降落值呈二次曲线相关关系 ;日温差与产量和千粒重呈二次曲线相关关系, 籽粒蛋白质和湿面筋含量、沉降值及降落值则随昼温差的增加线性递增 ;开花至成熟期降雨量与产量、千粒重都呈现先升后降的二次曲线相关关系, 而与籽粒蛋白质含量和降落值呈现线性负相关关系 ;籽粒蛋白质和湿面筋含量与降落值随开花至成熟期的累计日照时数呈现线性正相关关系。     

Comparative physiological response of wheat genotypes under terminal heat stress

Wheat (Triticum aestivum L.), a staple food crop, is of great commercial importance. Its production is restricted due to multiple environmental stresses. There are indications that the wheat production is consistently limited by terminal heat stress. Previous studies revealed a varied response of different wheat genotypes under heat stress conditions. Here, comparative physiological changes in wheat genotypes viz., DBW-140, Raj-3765, PBW-574, K-0-307 and HS-240 were evaluated under timely and late sown conditions in rabi season. We observed that heat stress dramatically affects chlorophyll content and leaf area index (LAI) in sensitive genotypes whereas proline and malondialdehyde (MDA) content were higher in tolerant genotypes under late sown conditions. Further, the heat susceptibility index (HIS) for 1,000-grain weight, grain weight and grain yield of wheat genotypes viz., HS 240 and K-0-307 was highest as compared with DBW 140, Raj 3765 and PBW 574 genotypes. This finding suggests that wheat genotypes are found to differ in their ability to respond to heat, thereby tolerance, which could be useful as genetic stock to develop wheat tolerant varieties in breeding programs.     

Gene action of canopy temperature in bread wheat under diverse environments

While canopy temperature (CT) shows a strong and reliable association with yield under drought and heat stress and is used in wheat breeding to select for yield, little is known of its genetic control. The objective of this study was to determine the gene action controlling CT in five wheat populations grown in diverse environments (heat, drought, and well-irrigated conditions). CT showed negative phenotypic correlations with grain yield under drought and well-irrigated environments. Additive × additive effects were most prevalent and significant for all crosses and environments. Dominance and dominance × dominance gene actions were also found, though the significance and direction was specific for each environment and genotypic cross. The use of CT as a selection criterion to improve tolerance to drought was supported by its significant association with grain yield and the genotype differences observed between cultivars. Our results indicated that genetic gains for CT in wheat could be achieved through conventional breeding. However, given some dominance and epistatic effects, it would be necessary to delay the selection process until the frequency of heterozygous loci within families is reduced.     

Quantitative trait loci for agronomic and physiological traits for a bread wheat population grown in environments with a range of salinity levels

Worldwide, salinity is a major environmental stress affecting agricultural production. Sodium (Na⁺) exclusion has long been recognised as a mechanism of salinity tolerance (ST) in cereals and several molecular markers have been suggested for breeding. However, there have been no empirical studies to show that selection for Na⁺ exclusion markers could improve grain yield in bread wheat under dryland salinity. In six field trials, a bread wheat mapping population was grown to validate Na⁺ exclusion quantitative trait loci (QTL) identified earlier in hydroponics, to determine the impact of Na⁺ exclusion on grain yield, and to identify QTL for yield-related traits. The traits included grain yield, grain number per m², 1,000-grain weight, maturity, plant height, and leaf Na⁺ and K⁺ concentrations. The presence of numerous QTL with minor effects for most traits indicated the genetic complexity of these traits, and thus limited prospects for pyramiding at present. Considerable QTL-by-environment interactions were observed, with the stable QTL generally being co-located with maturity or early vigour/height genes, which demonstrates the importance of measuring major agronomic traits in order to discover genuine QTL for ST. Several QTL for seedling biomass and Na⁺ exclusion identified earlier in hydroponics were also detected in field trials but with marginal impact on grain yield. These results suggest that selection for Na⁺ exclusion and the use of hydroponics-based seedling assays may not necessarily result in improved ST. However, as this is the first report of its kind, there is an urgent need for testing other mapping populations in realistic environments to discover novel ST-QTL for breeding programs. In the meantime, grain yield QTL independent of maturity and height may offer potential to improve ST.     

Enhanced N-metabolites,ABA and IAA-conjugate in anthers instigate heat sensitivity in spring wheat

Unraveling the metabolic and phytohormonal changes in anthers exposed to heat stress would help identify mechanisms regulating heat stress tolerance during the sensitive reproductive stage. Two spring wheat genotypes contrasting for heat tolerance were exposed to heat stress during heading in controlled environment chambers. Anthers were collected from main and primary spikes for metabolic and phytohormonal profiling. A significant reduction in seed set (38%), grain number (54%) and grain weight (52%) per plant was recorded in the sensitive (KSG1177) but not in the tolerant (KSG1214) genotype under heat stress compared to control. Anther metabolite accumulation did not vary quantitatively between main and primary spikes. Hierarchical clustering of the genotypes and treatments using metabolites and phytohormones revealed a distinct cluster for KSG1177 under heat stress from that of control and KSG1214. A significant increase in N-based amino acids, ABA, IAA-conjugate and a decrease in polyamines and organic acids were observed in wheat anthers exposed to heat stress. Unlike KSG1214, a significantly higher accumulation of amino acids, ABA and IAA-conjugate in anthers of the sensitive KSG1177 was recorded under heat stress. These findings provide the rationale and direction towards developing molecular markers for enhancing heat stress tolerance in wheat.     

小麦耐热性鉴定方法及热胁迫应答机理研究进展

小麦是全球最重要的粮食作物之一,高温特别是开花期和灌浆期的高温胁迫严重影响小麦的产量和品质。本文从直接鉴定法和间接鉴定法两个方面提出了小麦耐热性的评价指标;分析了在高温胁迫下温度信号的感知和传递以及植物对高温的响应机理;从传统育种方法和基因工程策略两个方向综述了小麦耐热性培育的研究进展。最后总结了提高小麦耐热性的措施及存在的问题,并提出了今后小麦耐热性研究展望。     

Single nucleotide polymorphism tightly linked to a major QTL on chromosome 7A for both kernel length and kernel weight in wheat

Thousand-kernel weight (TKW) is one of the major components of grain yield in wheat (Triticum aestivum). Identifying major quantitative trait loci (QTLs) for TKW and developing effective markers are prerequisite for success in marker-assisted selection (MAS) to improve wheat yield through breeding. This study mapped a major QTL, designated as TaTKW-7AL, for increasing TKW on the long arm of chromosome 7A of ‘Clark’ to a 1.3-cM interval between single nucleotide polymorphism (SNP) markers IWB13913 and IWA5913. This QTL explained 19.7 % of the phenotypic variation for TKW. A QTL for increasing kernel length (KL), one of the major components of TKW, was mapped in the same interval as TaTKW-7AL, suggesting that increased TKW by the QTL in ‘Clark’ is most likely due to the increased KL. Association analysis on a diversity panel of 200 US winter wheat accessions also identified a haplotype of three SNP markers (IWB13913, IWB6693 and IWA5913) that were tightly associated with the both KL and TKW. The analysis of allele frequencies of the haplotype in the diversity panel suggested that the favorable allele of TaTKW-7AL has not been strongly selected for in practice and has potential to be used to improve grain yield in US hard winter wheat breeding. Two user-friendly flanking KASPar markers, IWB13913 and IWA5913, were developed for MAS of TaTKW-7AL.     

Stay-green in spring wheat can be determined by spectral reflectance measurements (normalized difference vegetation index) independently from phenology

The green area displayed by a crop is a good indicator of its photosynthetic capacity, while chlorophyll retention or 'stay-green' is regarded as a key indicator of stress adaptation. Remote-sensing methods were tested to estimate these parameters in diverse wheat genotypes under different growing conditions. Two wheat populations (a diverse set of 294 advanced lines and a recombinant inbred line population of 169 sister lines derived from the cross between Seri and Babax) were grown in Mexico under three environments: drought, heat, and heat combined with drought. In the two populations studied here, a moderate heritable expression of stay-green was found-when the normalized difference vegetation index (NDVI) at physiological maturity was estimated using the regression of NDVI over time from the mid-stages of grain-filling to physiological maturity-and for the rate of senescence during the same period. Under heat and heat combined with drought environments, stay-green calculated as NDVI at physiological maturity and the rate of senescence, showed positive and negative correlations with yield, respectively. Moreover, stay-green calculated as an estimation of NDVI at physiological maturity and the rate of senescence regressed on degree days give an independent measurement of stay-green without the confounding effect of phenology. On average, in both populations under heat and heat combined with drought environments CTgf and stay-green variables accounted for around 30% of yield variability in multiple regression analysis. It is concluded that stay-green traits may provide cumulative effects, together with other traits, to improve adaptation under stress further.     

QTL mapping of terminal heat tolerance in hexaploid wheat (T. aestivum L.)

High temperature (>30 °C) at the time of grain filling is one of the major causes of yield reduction in wheat in many parts of the world, especially in tropical countries. To identify quantitative trait loci (QTL) for heat tolerance under terminal heat stress, a set of 148 recombinant inbred lines was developed by crossing a heat-tolerant hexaploid wheat (Triticum aestivum L.) cultivar (NW1014) and a heat-susceptible (HUW468) cultivar. The F(5), F(6), and F(7) generations were evaluated in two different sowing dates under field conditions for 2 years. Using the trait values from controlled and stressed trials, four different traits (1) heat susceptibility index (HSI) of thousand grain weight (HSITGW); (2) HSI of grain fill duration (HSIGFD); (3) HSI of grain yield (HSIYLD); and (4) canopy temperature depression (CTD) were used to determine heat tolerance. Days to maturity was also investigated. A linkage map comprising 160 simple sequence repeat markers was prepared covering the whole genome of wheat. Using composite interval mapping, significant genomic regions on 2B, 7B and 7D were found to be associated with heat tolerance. Of these, two (2B and 7B) were co-localized QTL and explained more than 15 % phenotypic variation for HSITGW, HSIGFD and CTD. In pooled analysis over three trials, QTL explained phenotypic variation ranging from 9.78 to 20.34 %. No QTL × trial interaction was detected for the identified QTL. The three major QTL obtained can be used in marker-assisted selection for heat stress in wheat.     

Heat susceptibility of grain filling in wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) linked with rapid chlorophyll loss during a 3-day heat treatment

Brief heat events (1–3 days, >30 °C) commonly reduce wheat (Triticum aestivum L.) grain size and consequently yield. To identify mechanisms of tolerance to such short heat events, 36 wheat genotypes were treated under day/night temperatures of 37 °C/27 °C for 3-days in a growth chamber, at 10 days after anthesis, and a range of developmental, chlorophyll and yield-related traits monitored. The degree of flag leaf chlorophyll loss during the treatment was the variable that showed the highest correlation to grain weight loss (r = 0.63; p < 0.001), identifying chlorophyll stability during this brief period as a potential determinant or indicator of grain weight stability under heat. Variables summarizing the combined during- and post-heat chlorophyll losses showed similar or lower correlations with heat tolerance of grain filling, despite the fact that genotypes varied in their ability to resume normal chlorophyll loss rates after the heat treatment. Additionally, heat tolerance of grain size showed no correlation with grain filling duration or traits relating to utilization of stem carbon reserves under heat stress. Measurement of chlorophyll loss over a forecasted heat wave was thereby identified as a potential basis for developing tools to help breeders select heat tolerant genotypes.     

Breeding Enrichment of Genetic Variation of Grain Yield and Its Attributes in Bread Wheat under Drought Stress and Well Irrigation

Drought stress (DS) is one of the most critical environmental abiotic stresses for wheat production in the arid environments. Selection of high-yielding genotypes tolerant to DS can play a significant role in mitigation the negative impacts associated with DS. In the present study, generation means analysis (GMA) was used to study the performance of two crosses under well irrigation (WI) and deficit irrigation [cross I (Line 44 × Shandweel-1) and cross II (Line 20 × Sakha 93)]. Significant differences were observed for days to heading (DH), days to maturity (DM), plant height (PH), spike length (SL), number of spikes per plant (NS/P), number of grains per spike (NG/S), thousand-grain weight (TGW), grain yield per plant (GY/P), and proline content (PC) in the six populations of the two crosses within each irrigation level. Cross II had early maturity and the highest PC, NS/P, TGW, and GY/P regardless of the irrigation level. Cross I showed positive significant relative heterosis and heterobeltiosis for GY/P under the two irrigation levels. The inheritance of characters of cross I revealed additive, dominant, and epistatic effects, which varied with trait and stress. Additive genetic effects predominated in DH, SL, and PC, while non- additive were found in DM, NS/P, NG/S, and GY/P. Narrow-sense heritability estimates (h²_n) were high for DH and PC, moderate to high for PH and SL, moderate for DM, NG/S, NS/P, and TGW, and low for GY/P. Based on different drought indices the populations BC₁, BC₂, F₁, and P₁ of cross II and BC1 of cross I were more tolerant to drought stress. Therefore, PC, TGW and DH can be used as selection indicators to improve wheat for drought tolerance in early generations and other yield components traits in late generations. The second cross (Line 20 × Sakha 93) shows promise and is of interest to a drought tolerance breeding program, where wheat breeders can use recombinant breeding strategies to construct desirable drought stress genes. Correlation and path coefficient revealed that TGW and PC were the main contributor in grain yield in both environments.     

Proteomics uncovers a role for redox in drought tolerance in wheat

Proteomic analysis offers a new approach to identify a broad spectrum of genes that are expressed in living systems. We applied a proteomic approach to study changes in wheat grain in response to drought, a major environmental parameter adversely affecting development and crop yield. Three wheat genotypes differing in genetic background were cultivated in field under well-watered and drought conditions by following a randomized complete block design with four replications. The overall effect of drought was highly significant as determined by grain yield and total dry matter. About 650 spots were reproducibly detected and analyzed on 2-DE gels. Of these, 121 proteins showed significant change under drought condition in at least one of the genotypes. Mass spectrometry analysis using MALDI-TOF/TOF led to the identification of 57 proteins. Two-thirds of identified proteins were thioredoxin (Trx) targets, in accordance with the link between drought and oxidative stress. Further, because of contrasting changes in the tolerant and susceptible genotypes studied, several proteins emerge as key participants in the drought response. In addition to providing new information on the response to water deprivation, the present study offers opportunities to pursue the breeding of wheat with enhanced drought tolerance using identified candidate genetic markers. The 2-DE database of wheat seed proteins is available for public access at http://www.proteome.ir.     

Genetic Variation and Association Analysis of the SSR Markers Linked to the Major Drought-Yield QTLs of Rice

Drought is one of the major abiotic stresses, which hampers the production of rice worldwide. Informative molecular markers are valuable tools for improving the drought tolerance in various varieties of rice. The present study was conducted to evaluate the informative simple sequence repeat (SSR) markers in a diverse set of rice genotypes. The genetic diversity analyses of the 83 studied rice genotypes were performed using 34 SSR markers closely linked to the major quantitative trait loci (QTLs) of grain yield under drought stress (qDTYs). In general, our results indicated high levels of polymorphism. In addition, we screened these rice genotypes at the reproductive stage under both drought stress and nonstressful conditions. The results of the regression analysis demonstrated a significant relationship between 11 SSR marker alleles and the plant paddy weight under stressful conditions. Under the nonstressful conditions, 16 SSR marker alleles showed a significant correlation with the plant paddy weight. Finally, four markers (RM279, RM231, RM166, and RM231) demonstrated a significant association with the plant paddy weight under both stressful and nonstressful conditions. These informative-associated alleles may be useful for improving the crop yield under both drought stress and nonstressful conditions in breeding programs.     

Evaluation of Wheat Chromosome Translocation Lines for High Temperature Stress Tolerance at Grain Filling Stage

High temperature (HT, heat) stress is detrimental to wheat (Triticum aestivum L.) production. Wild relatives of bread wheat may offer sources of HT stress tolerance genes because they grow in stressed habitats. Wheat chromosome translocation lines, produced by introgressing small segments of chromosome from wild relatives to bread wheat, were evaluated for tolerance to HT stress during the grain filling stage. Sixteen translocation lines and four wheat cultivars were grown at optimum temperature (OT) of 22/14°C (day/night). Ten days after anthesis, half of the plants were exposed to HT stress of 34/26°C for 16 d, and other half remained at OT. Results showed that HT stress decreased grain yield by 43% compared with OT. Decrease in individual grain weight (by 44%) was the main reason for yield decline at HT. High temperature stress had adverse effects on leaf chlorophyll content and Fv/Fm; and a significant decrease in Fv/Fm was associated with a decline in individual grain weight. Based on the heat response (heat susceptibility indices, HSIs) of physiological and yield traits to each other and to yield HSI, TA5594, TA5617, and TA5088 were highly tolerant and TA5637 and TA5640 were highly susceptible to HT stress. Our results suggest that change in Fv/Fm is a highly useful trait in screening genotypes for HT stress tolerance. This study showed that there is genetic variability among wheat chromosome translocation lines for HT stress tolerance at the grain filling stage and we suggest further screening of a larger set of translocation lines.     

Heat Stress-Induced Cell Death, Changes in Antioxidants, Lipid Peroxidation, and Protease Activity in Wheat Leaves

Six wheat genotypes were evaluated for heat tolerance in terms of seedling growth, antioxidant response and cell death. Based on the heat susceptibility index (HSI), response of the genotypes varied from heat tolerant (Inqilab-91) to heat sensitive (Sitta) along with moderately tolerant (Nesser and Sarsabz) and sensitive (Fareed and FD-83). Heat stress-induced programmed cell death (probably necrosis) in wheat leaves was evident by DNA smear. MDA content increased above twofold in most of genotypes under heat stress, with the lowest increase in the heat-tolerant genotype Nesser. Catalase activity diminished under heat stress in all genotypes. Peroxidase, superoxide dismutase (SOD), protease, and ascorbate peroxidase (APX) activities increased under heat stress. Apparently, heat stress-induced reduction in catalase activity was compensated by a parallel increase in peroxidases to quench H₂O₂. Heat stress-induced decrease (%) in catalase and increase in protease activities showed significant positive correlations, whereas increase (%) in APX activity showed a significant negative correlation with HSI or relative heat tolerance of genotypes. All these correlations signify that catalase, protease and ascorbate peroxidase can be used efficiently as biochemical markers to assess the relative heat stress tolerance of wheat genotypes at the seedling stage. In conclusion, using a multiparametric approach involving morphophysiological and biochemical assays, the sensitivity of wheat genotypes to heat stress could be evaluated to a sufficient level of certainty at the seedling stage.     

Characterization of some bread wheat genotypes using molecular markers for drought tolerance

Because of its wide geographical adaptation and importance in human nutrition, wheat is one of the most important crops in the world. However, wheat yield has reduced due to drought stress posing threat to sustainability and world food security in agricultural production. The first stage of drought tolerant variety breeding occurs on the molecular and biochemical characterization and classification of wheat genotypes. The aim of the present study is characterization of widely grown bread wheat cultivars and breeding lines for drought tolerance so as to be adapted to different regions in Turkey. The genotypes were screened with molecular markers for the presence of QTLs mapped to different chromosomes. Results of the molecular studies identified and detected 15 polymorphic SSR markers which gave the clearest PCR bands among the control genotypes. At the end of the research, bread wheat genotypes which were classified for tolerance or sensitivity to drought and the genetic similarity within control varieties were determined by molecular markers. According to SSR based dendrogram, two main groups were obtained for drought tolerance. At end of the molecular screening with SSR primers, genetic similarity coefficients were obtained that ranged from 0.14 to 0.71. The ones numbered 8 and 11 were the closest genotypes to drought tolerant cultivar Gerek 79 and the furthest genotypes from this cultivar were number 16 and to drought sensitive cultivar Sultan 95. The genotypes as drought tolerance due to their SSR markers scores are expected to provide useful information for drought related molecular breeding studies.     

Wheat cultivars differing in heat tolerance show a differential response to monocarpic senescence under high-temperature stress and the involvement of serine proteases

High temperature is a common constraint during anthesis and grain-filling stages of wheat leading to huge losses in yield. In order to understand the mechanism of heat tolerance during monocarpic senescence, the present study was carried out under field conditions by allowing two well characterized Triticum aestivum L. cultivars differing in heat tolerance, Hindi62 (heat-tolerant) and PBW343 (heat-susceptible), to suffer maximum heat stress under late sown conditions. Senescence was characterized by measuring photosynthesis related processes and endoproteolytic activity during non-stress environment (NSE) as well as heat-stress environment (HSE). There was a faster rate of senescence under HSE in both the genotypes. Hindi62, having pale yellow flag leaf with larger area, maintained cooler canopy under high temperatures than PBW343. The tolerance for high temperature in Hindi62 was clearly evident in terms of slower green-leaf area degradation, higher stomatal conductance, higher stability in maximum PSII efficiency, Rubisco activity and Rubisco content than PBW343. Both the genotypes exhibited lower endopeptidase activity under HSE as compared to NSE and this difference was more apparent in Hindi62. Serine proteases are the predominant proteases responsible for protein degradation under NSE as well as HSE. Flag leaf of both the genotypes exhibited high-molecular-mass endoproteases (78 kDa and 67 kDa) isoforms up to full grain maturity which were inhibited by specific serine protease inhibitor in both the environments. In conclusion, the heat-tolerant Hindi62 exhibited a slower rate of senescence than the heat-susceptible PBW343 during HSE, which may contribute towards heat stability.