Heat tolerance in spring wheat. I. Estimating cellular thermotolerance and its heritability

High temperature is a major environmental stress factor limiting wheat (Triticum aestivum L.) productivity. Improvement of heat tolerance in wheat is an important breeding objective. Genetic variation in cellular thermotolerance among 56 spring wheat cultivars was evaluated at the seedling stage of growth by cell membrane thermostability (CMS) and triphenyl tetrazolium chloride (TTC) assays. A subset of eight lines was also evaluated at the flowering stage using the same assays. With both assays Average thermotolerance tended to decrease from the seedling to the flowering stages. However, thermotolerance was well correlated between growth stages among the eight cultivars for both CMS (r = 0.92; p = 0.004) and TTC (r = 0.84; p = 0.050). The correlation between TTC and CMS among the eight cultivars tested at the seedling and the flowering growth stages was significant (r = 0.74; p = 0.031 and r = 0.75; p = 0.029, respectively). The same correlation was less strong, though still significant (r = 0.32; p = 0.014) across 56 cultivars at the seedling stage. In a study of the cross V747 (tolerant)/Barkaee (susceptible), broad sense heritability was estimated at 89% for TTC. Most of the genetic variance was additive. CMS in seedlings of 16 cultivars was positively and significantly (p ≤ 0.05) correlated with yields of these cultivars in each of four hot environments in Mexico, Sudan, India, and Brazil. The same correlations for TTC were positive but nonsignificant. This revised version was published online in August 2006 with corrections to the Cover Date.     

Maize mesocotyl: Role in response to stress and deep-sowing tolerance

Maize (Zea mays L.) is one of the most important crops worldwide and is a model organism among cereal crops. Abiotic and biotic stresses are often present simultaneously and severely influence maize production, causing great yield losses worldwide. Therefore, the selection and cultivation of stress-tolerant maize lines that adapt to various stresses is instrumental in addressing the problem of yield losses caused by stress. The maize mesocotyl is the crucial organ that pushes shoots out of deep water or soil after seed germination. It has a simple anatomy and exhibits rapid growth in the dark. In this article, we reviewed the studies on the elongation of the maize mesocotyl and the actions of phytohormones, especially under deep-sowing conditions, and emphasized the role of the maize mesocotyl in response to environmental stress and deep-sowing tolerance. We propose that the maize mesocotyl can serve as a selection organ for evaluating stress tolerance at the early seedling stage. We also identify future research fields that need further investigation in studies of the maize mesocotyl.     

Integrated breeding approaches for improving drought and heat adaptation in chickpea (Cicer arietinum L.)

Chickpea (Cicer arietinum L.) is a dry season food legume largely grown on residual soil moisture after the rainy season. The crop often experiences moisture stress towards end of the crop season (terminal drought). The crop may also face heat stress at the reproductive stage if sowing is delayed. The breeding approaches for improving adaptation to these stresses include the development of varieties with early maturity and enhanced abiotic stress tolerance. Several varieties with improved drought tolerance have been developed by selecting for grain yield under moisture stress conditions. Similarly, selection for pod set in the crop subjected to heat stress during reproductive stage has helped in the development of heat‐tolerant varieties. A genomic region, called QTL‐hotspot, controlling several drought tolerance‐related traits has been introgressed into several popular cultivars using marker‐assisted backcrossing (MABC), and introgression lines giving significantly higher yield than the popular cultivars have been identified. Multiparent advanced generation intercross (MAGIC) approach has been found promising in enhancing genetic recombination and developing lines with enhanced tolerance to terminal drought and heat stresses.     

Heat,wheat and CO2: The relevance of timing and the mode of temperature stress on biomass and yield

Atmospheric CO₂ enrichment affects C3 crops both directly via increased carbon gain and improved water use efficiency and indirectly via higher temperatures and more frequent climatic extremes. Here we investigated the response of spring wheat (Triticum aestivum L. cv. Triso) to CO₂ enrichment (550 vs. 380 µmol/mol) and heat, applied as a constant +4°C increase or a typical heat wave either before or after anthesis, or as two typical heat waves before and after anthesis. We applied a climate chamber approach closely mimicking ambient conditions. CO₂ enrichment increased above‐ground biomass and yield by c. 7 and 10%, but was not able to compensate for adverse heat stress effects, neither before nor after anthesis, with few exceptions only. Yield depression due to heat stress was most severe when two heat waves were applied (?19%). This adverse effect was, however, compensated by CO₂ enrichment. Applying heat stress before or after anthesis did not exert different effects on yield for both +4°C warming and heat wave application. However, +4°C depressed yield more than a heat wave at ambient CO₂, but not so at elevated CO₂. Thus, the interactive effects were complex and prediction of future wheat yield under CO₂ enrichment and climate extremes deserves more attention.     

Cowpea (Vigna unguiculata): Genetics,genomics and breeding

Cowpea, Vigna unguiculata (L.), is an important grain legume grown in the tropics where it constitutes a valuable source of protein in the diets of millions of people. Some abiotic and biotic stresses adversely affect its productivity. A review of the genetics, genomics and breeding of cowpea is presented in this article. Cowpea breeding programmes have studied intensively qualitative and quantitative genetics of the crop to better enhance its improvement. A number of initiatives including Tropical Legumes projects have contributed to the development of cowpea genomic resources. Recent progress in the development of consensus genetic map containing 37,372 SNPs mapped to 3,280 bins will strengthen cowpea trait discovery pipeline. Several informative markers associated with quantitative trait loci (QTL) related to desirable attributes of cowpea were generated. Cowpea genetic improvement activities aim at the development of drought tolerant, phosphorus use efficient, bacterial blight and virus resistant lines through exploiting available genetic resources as well as deployment of modern breeding tools that will enhance genetic gain when grown by sub‐Saharan Africa farmers.     

Field‐grown Cotton Exhibits Seasonal Variation in Photosynthetic Heat Tolerance without Exposure to Heat‐stress or Water‐deficit Conditions

Thermotolerance acclimation of photosystem II to heat and drought is well documented, but studies demonstrating developmental impacts on heat tolerance in field‐grown plants are limited. Consequently, climatic variables, estimated canopy temperature, predawn leaf water potential (Ψ_PD), and the temperature responses of maximum quantum yield of photosystem II (F_v/F_m), variable fluorescence (F_v/F₀), quantum yield of electron transport (φ_Eο) and efficiency of PSI electron acceptor reduction (REο/ABS) were characterized for Gossypium hirsutum at three sample times during the growing season (21 June, 2 July and 18 July 2013) under well‐watered conditions. The temperature decreasing a given photosynthetic parameter 15% from the optimum is referred to as T₁₅ and served as a standardized measure of heat tolerance. Ambient and estimated canopy temperatures were well within the optimal range for cotton throughout the sample period, and leaves were verified well watered using Ψ_PD measurements. However, T₁₅ varied with sample date (highest on July 2 for all parameters), being 2 °C (F_v/F₀) to 5.5 °C (φ_Eο) higher on July 2 relative to June 21, despite optimal temperature conditions and predawn leaf water potential on all sample dates. These findings suggest that even under optimum temperature conditions and water availability, heat tolerance could be influenced by plant developmental stage.     

Crop breeding for salt tolerance in the era of molecular markers and marker‐assisted selection

Crop salt tolerance (ST) is a complex trait affected by numerous genetic and non‐genetic factors, and its improvement via conventional breeding has been slow. Recent advancements in biotechnology have led to the development of more efficient selection tools to substitute phenotype‐based selection systems. Molecular markers associated with genes or quantitative trait loci (QTLs) affecting important traits are identified, which could be used as indirect selection criteria to improve breeding efficiency via marker‐assisted selection (MAS). While the use of MAS for manipulating simple traits has been streamlined in many plant breeding programmes, MAS for improving complex traits seems to be at infancy stage. Numerous QTLs have been reported for ST in different crop species; however, few commercial cultivars or breeding lines with improved ST have been developed via MAS. We review genes and QTLs identified with positive effects on ST in different plant species and discuss the prospects for developing crop ST via MAS. With the current advances in marker technology and a better handling of genotype by environment interaction effects, the utility of MAS for breeding for ST will gain momentum.     

Heat tolerance for yield and its components in different wheat cultivars

Summary Twenty one diverse, standard and experimental cultivars of common spring wheat (Triticum aestivum L.) were tested for the effect of heat stress on phenology, yield and its components by growing the materials for 2 years under full irrigation during the hot summer (offseason), and the cool winter (normal) conditions. Heat tolerance was estimated for each variable by the heat susceptibility index (S) which scales the reduction in cultivar performance from cool to hot conditions relative to the respective mean reduction over all cultivars.Genotypes differed significantly in S for yield and its components. The ranking of cultivars in S over the 2 years was consistent for yield, kernels per spike and kernel weight, but not for spike number. Of the three yield components, the greatest genotypic variation in S was expressed for kernels per spike. However, S for yield could not be simply attributed to S in a unique component across all cultivars. On the other hand, a general linear model regression of summer yield on its components revealed that the most important yield component affecting yield variation among cultivars under heat stress was kernel number per spike. Kernel number per spike was positively associated across cultivars with longer duration and greater stabilty of thermal time requirement from emergence to double ridge. It is therefore concluded that kernel number per spike under heat stress is a reasonable estimate of heat tolerance in yield of wheat and that this tolerance is operative already during the first 2 to 3 weeks of growth.     

Impact of wheat-Leymus racemosus added chromosomes on wheat adaptation and tolerance to heat stress

Adaptation of wheat (Triticum aestivum L.) to high temperatures could be improved by introducing alien genes from wild relatives. We evaluated the responses of wheat-Leymus racemosus chromosome introgression lines to high temperature to determine their potentiality for developing improved wheat cultivars. Introgression lines and their parent Chinese Spring were evaluated in a growth chamber at the seedling stage and in the field at the reproductive stage in two heat-stressed environments in Sudan. Optimum and late planting were used to ensure exposure of the plants to heat stress at the reproductive stage. The results revealed the impact of several Leymus chromosomes in improving wheat adaptation and tolerance to heat. Three lines possessed enhanced adaptation, whereas two showed high heat tolerance. Two addition lines showed a large number of kernels per spike, while one possessed high yield potential. Grain yield was correlated negatively with the heat susceptibility index, days to heading and maturity and positively with kernel number per spike and triphenyl tetrazolium chloride assay under late planting. The findings suggest that these genetic stocks could be used as a bridge to introduce the valuable Leymus traits into a superior wheat genetic background, thus helping maximize wheat yield in heat-stressed environments.     

Genetic diversity of a global collection of maize genetic resources in relation to their subspecies assignments,geographic origin,and drought tolerance

The genetic diversity among an international collection of 40 maize accessions has been evaluated using DNA ISSR fingerprinting. Among the 180 ISSR markers scored by 15 primers, 161 markers (89.59%) were polymorphic and 19 were unique in 16 accessions. A cluster tree based on the average distance coefficients and the Dice similarity indices divided the accessions into three major groups, each including clusters of accessions assigned to their subspecies. However, a low level of genetic differentiation among the accessions was demonstrated by the STRUCTURE analysis of ISSR data in agreement with the low gene flow (Nm) value among the accessions. A scatter diagram of the principal component analysis (PCA) based on ISSR data analysis revealed that the accessions were differentiated into three groups comparable to those produced by the cluster analysis, in which some accessions of the same subspecies showed a close similarity to each other. A scatter diagram of the principal coordinate analysis (PCoA) based on the drought tolerance indices (DTIs) showed that nine genetically similar accessions share drought tolerance characteristics; these include four of subsp. indurata, three of subsp. everata, and two of subsp. indentata. An abundance of unique ISSR alleles found in the 16 accessions, including the nine drought-tolerant accessions, represents rich untapped genetic resources and these accessions may be exploited in the future breeding of maize commercial lines.     

Breeding for boron tolerance in lentil (Lens culinaris Medik.) using a high‐throughput phenotypic assay and molecular markers

This study describes the identification of a quantitative trait locus (QTL) in the recombinant inbred line population of ILL2024 × ILL6788 and subsequent validation of associated molecular markers. A high‐quality genetic linkage map was constructed with 758 markers that cover 1,057 cM, with an average intermarker distance of 2 cM. QTL analysis revealed a single genomic region on Lc2 to be associated with B tolerance and accounted for up to 76% of phenotypic variation (Vp). The best markers for B tolerance were assessed for their utility in routine breeding applications using validation panels of diverse lentil germplasm and breeding material derived from ILL2024. A marker generated from the dense genetic map of this study was found to be the most accurate of all markers available for B tolerance in lentil, with a success rate of 93% within a large breeding pool derived from ILL2024. However, given the number of the unrelated lines for which the marker–trait association was not conserved, B tolerance screening is still required at later stages to confirm predicted phenotypes.     

Susceptibility of Faba Bean (Vicia faba L.) to Heat Stress During Floral Development and Anthesis

Experiments were conducted over 2 years to quantify the response of faba bean (Vicia faba L.) to heat stress. Potted winter faba bean plants (cv. Wizard) were exposed to temperature treatments (18/10; 22/14; 26/18; 30/22; 34/26 °C day/night) for 5 days during floral development and anthesis. Developmental stages of all flowers were scored prior to stress, plants were grown in exclusion from insect pollinators to prevent pollen movement between flowers, and yield was harvested at an individual pod scale, enabling effects of heat stress to be investigated at a high resolution. Susceptibility to stress differed between floral stages; flowers were most affected during initial green‐bud stages. Yield and pollen germination of flowers present before stress showed threshold relationships to stress, with lethal temperatures (t₅₀) ?28 °C and ~32 °C, while whole plant yield showed a linear negative relationship to stress with high plasticity in yield allocation, such that yield lost at lower nodes was partially compensated at higher nodal positions. Faba bean has many beneficial attributes for sustainable modern cropping systems but these results suggest that yield will be limited by projected climate change, necessitating the development of heat tolerant cultivars, or improved resilience by other mechanisms such as earlier flowering times.     

Cold stress tolerance of soybeans during flowering: QTL mapping and efficient selection strategies under controlled conditions

Breeding soybeans for higher latitudes requires cultivars with an increased chilling stress tolerance, especially when flowering occurs. Phenotyping in climate chambers to select for this trait is labour‐intensive and requires an optimal allocation of resources due to limited space. We screened a diversity panel of 35 early maturity cultivars and a biparental population of 103 RILs for their cold stress tolerance at flowering stage. Pod number under control and stress conditions is highly heritable and showed only a weak correlation between the two treatments. Based on different testing scenarios, we could show that testing more genotypes with less replicates yields much higher responses to selection and hence should be pursued in such climate‐controlled experiments. We identified quantitative trait loci (QTL) for pod number under both conditions (chromosomes 7 and 13) and a cold tolerance‐specific QTL (chromosome 11). Furthermore, we performed genomic predictions using different test set scenarios and prediction models, showing that genomic prediction is a promising tool to select for cold stress tolerance, particularly if known QTL can be used as fixed effects in the model.     

Evaluation of heat stress tolerance in irrigated environment of T. aestivum and related species. I. Stability in yield and yield components

Wheat production is often limited by continual or terminal heat stress. The current study was aimed at the characterization of wild relatives and cultivated Triticum species for their heat tolerance in yield and its analysis in relation to yield components which confer yield stability at the three ploidy levels. Thirty-two non-cultivated and cultivated genotypes belonging to diploid, tetraploid and hexaploid wheat species were evaluated for heat stress tolerance in the field under full irrigation. Wheat species were sown in the field(New Delhi, India; 77°12′E, 28°40′N, 228.6 m m.s.l) at two dates of sowing, November (normal) and January (late Sowing) during winter seasons of 1994–95 and 1995–96. The late sown crop experienced 3°C warmer temperatures than that of the normal sown crop. Wide variability was observed for grain yield stability under heat stress, as the heat susceptibility index (S) ranged from 0.13 to 2.08. Hexaploidy conferred the productive and adaptive advantages as it combined high yield and stability when compared to the tetraploid and diploid groups. However within each ploidy group wide variation was observed for heat tolerance. T. aestivum cv C306 & HI1136, T. dicoccoides, T. monococcum acc. BSP1 and Ae. speltoides ssp. liqustica were highly heat tolerant in their grain yield. Stability in grain no. m- 2 conferred yield stability in all three ploidy levels, although grain weight stability also contributed to yield stability in moderately stable T. turgidum and T. sphaerococcum under heat stress. Higher biomass and grain no. m-2 are the two important traits which could be considered potential selection criteria for yield under heat stress. Of the two components of grain no. m-2, stability in spike no. m-2could be considered more important trait than grain no. spike-1. Since wide variation for heat tolerance of all the yield components are available among the wheat species, these species can be used for improving specific yield components of cultivated wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Use of barley seed vigour to discriminate drought and cold tolerance in crop years with high seed vigour and low trait variation

Seed vigour is a precondition for early and homogenous field emergence of barley, in addition to effective malting. This study aimed to assess the selection of barley varieties by using seed vigour as the indicator. Seed vigour of barley (quantified as the germination percentage) was evaluated under drought (?0.2 MPa) and temperature stress (10°C). At two locations over a 3‐year period, 1 population of 133 Derkado × B83‐12/21/5 doubled haploid (DH) lines (and parents) was evaluated for seed vigour, of which 108 DH lines were assessed for three malting parameters. The relatively high values of vigour during the 3‐year period (overall average 94–95%) probably impeded high variations in genetic potential. A total of 27 DH lines of the 133 evaluated showed transgression for vigour (up to 98%) in comparison with the parents (Derkado: 96%; B83: 92%). In conclusion, caution should be applied when selecting for seed vigour, even in good crop years with high levels of seed vigour and low trait variations. Such selection might improve vigour, particularly in crop years with unsuitable weather conditions.     

Comparison of QTLs for seed germination under non-stress, cold stress and salt stress in tomato

The purpose of this study was to examine whether rate of tomato seed germination under non-stress, cold-stress and salt-stress conditions was under similar genetic control by identifying and comparing quantitative trait loci (QTLs) which affect germination rate under these conditions. A fast-germinating accession (LA722) of the wild tomato species Lycopersicon pimpinellifolium Jusl. and a slow-germinating cultivar (NC84173, maternal and recurrent parent) of tomato (Lycopersicon esculentum Mill.) were hybridized and BC₁ and BC₁S₁ progeny produced. The BC₁ population was used to construct a linkage map with 151 restriction fragment length polymorphism (RFLP) markers. The BC₁S₁ population (consisting of 119 BC₁S₁ families) was evaluated for germination under non-stress (control), cold-stress and salt-stress conditions and the mean time to 50% germination (T50) in each treatment was determined. Germination analyses indicated the presence of significant (P < 0.01) phenotypic correlations between T50 under control and cold stress (r = 0.71), control and salt stress (r = 0.58) and cold stress and salt stress (r = 0.67). The QTL analysis indicated the presence of genetic relationships between germination under these three conditions: a few QTLs were identified which commonly affected germination under both stress- (cold-, salt- or both) and non-stress conditions, and thus were called stress-nonspecific QTLs. A few QTLs were also identified which affected germination only under cold or salt stress and thus were called stress-specific QTLs. However, the stress-nonspecific QTLs generally exhibited larger individual effects and together accounted for a greater portion of the total phenotypic variation under each condition than the stress-specific QTLs. Whether the effects of stress-nonspecific QTLs were due to pleiotropic effects of the same genes, physical linkage of different genes, or a combination of both could not be determined in this study. The results, however, indicate that the rate of tomato seed germination under different stress and nonstress conditions is partly under the same genetic control.     

Characterization of a set of common wheat–Hordeum chilense chromosome 7Hch introgression lines and its potential use in research on grain quality traits

Chromosome 7H^ch from Hordeum chilense has potential for improving seed carotenoid content in wheat as it carries a Phytoene synthase 1 (Psy1) gene, which has a major role in this trait. Structural changes in chromosome 7H^ch were obtained in common wheat background by crossing the wheat disomic substitution line 7H^ch(7D) with a disomic addition line carrying chromosome 2C^c from Aegilops cylindrica in common wheat cv. ‘Chinese Spring’. Rearranged 7H^ch chromosomes were cytologically characterized by FISH. A set of 24 molecular markers and the Psy1 gene were used to identify the H. chilense chromosome segments involved in the introgressions. Six structural rearrangements of chromosome 7H^ch were identified. They included three homozygous wheat–H. chilense centromeric translocations, one involving the 7H^chS arm (T‐7H^chS·A/B) and two involving the 7H^chL arm (T1‐7H^chL·A/B and T2‐7H^chL·A/B). In addition, one 7H^chS arm deletion, one 7H^chL·7H^chL isochromosome and one 7H^chS telosome were obtained in hemizygous condition. These genetic stocks will be useful for studying the effect of chromosome 7H^ch on wheat flour colour.     

Effect of high temperature on pollen morphology,plant growth and seed yield in quinoa (Chenopodium quinoa Willd.)

Quinoa (Chenopodium quinoa Willd.) has gained considerable attention worldwide during the past decade due to its nutritional and health benefits. However, its susceptibility to high temperatures has been reported as a serious obstacle to its global production. The objective of this study was to evaluate quinoa growth and pollen morphology in response to high temperatures. Pollen morphology and viability, plant growth and seed set, and several physiological parameters were measured at anthesis in two genotypes of quinoa subjected to day/night temperatures of 22/16°C as a control treatment and 40/24°C as the heat stress treatment. Our results showed that heat stress reduced the pollen viability between 30% and 70%. Although no visible morphological differences were observed on the surface of the pollen between the heat‐stressed and non‐heat‐stressed treatments, the pollen wall (intine and extine) thickness increased due to heat stress. High temperature did not affect seed yield, seed size and leaf greenness. On the other hand, high temperature improved the rate of photosynthesis. We found that quinoa has a high plasticity in response to high temperature, though pollen viability and pollen wall structure were affected by high temperatures in anthesis stage. This study is also the first report of quinoa pollen being trinucleate.     

Current status and future strategy in breeding chickpea for resistance to biotic and abiotic stresses

Chickpea (Cicer arietinum L.) production has remained static for the past two decades. One major limiting factor has been susceptibility of cultivars to several biotic and abiotic stresses that adversely affect yield. In recent years, cultivars resistant to Ascochyta blight (Ascochyta rabiei [Pass.] Lab.), Fusarium wilt (Fusarium oxysporum f. sp. ciceris), and cold have been bred and released in many countries. Some progress has been made in breeding for resistance to drought, insects, and cyst nematode, but not for viruses, heat, and salinity. Two or more stresses are of equal importance in most chickpea growing areas. Therefore, future efforts should be directed toward the development of cultivars with multiple-stress resistance. Proper understanding of important stresses in different countries and the genetics of resistance should lead to more systematic approaches to resistance breeding. Wild Cicer species hold promise and deserve attention in resistance breeding.