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.     

Abiotic stresses,constraints and improvement strategies in chickpea

Chickpea (Cicer arietinum L.) is cultivated mostly in the arid and semi‐arid regions of the world. Climate change will bring new production scenarios as the entire growing area in Indo–Pak subcontinent, major producing area of chickpea, is expected to undergo ecological change, warranting strategic planning for crop breeding and husbandry. Conventional breeding has produced several high‐yielding chickpea genotypes without exploiting its potential yield owing to a number of constraints. Among these, abiotic stresses include drought, salinity, water logging, high temperature and chilling frequently limit growth and productivity of chickpea. The genetic complexity of these abiotic stresses and lack of proper screening techniques and phenotyping techniques and genotype‐by‐environment interaction have further jeopardized the breeding programme of chickpea. Therefore, considering all dispiriting aspects of abiotic stresses, the scientists have to understand the knowledge gap involving the physiological, biochemical and molecular complex network of abiotic stresses mechanism. Above all emerging ‘omics’ approaches will lead the breeders to mine the ‘treasuring genes’ from wild donors and tailor a genotype harbouring ‘climate resilient’ genes to mitigate the challenges in chickpea production.     

Chickpea molecular breeding: New tools and concepts

Summary Chickpea is a cool season grain legume of exceptionally high nutritive value and most versatile food use. It is mostly grown under rain fed conditions in arid and semi-arid areas around the world. Despite growing demand and high yield potential, chickpea yield is unstable and productivity is stagnant at unacceptably low levels. Major yield increases could be achieved by development and use of cultivars that resist/tolerate abiotic and biotic stresses. In recent years the wide use of early maturing cultivars that escape drought stress led to significant increases in chickpea productivity. In the Mediterranean region, yield could be increased by shifting the sowing date from spring to winter. However, this is hampered by the sensitivity of the crop to low temperatures and the fungal pathogen Ascochyta rabiei. Drought, pod borer (Helicoverpa spp.) and the fungus Fusarium oxysporum additionally reduce harvests there and in other parts of the world. Tolerance to rising salinity will be a future advantage in many regions. Therefore, chickpea breeding focuses on increasing yield by pyramiding genes for resistance/tolerance to the fungi, to pod borer, salinity, cold and drought into elite germplasm. Progress in breeding necessitates a better understanding of the genetics underlying these traits. Marker-assisted selection (MAS) would allow a better targeting of the desired genes. Genetic mapping in chickpea, for a long time hampered by the little variability in chickpea’s genome, is today facilitated by highly polymorphic, co-dominant microsatellite-based markers. Their application for the genetic mapping of traits led to inter-laboratory comparable maps. This paper reviews the current situation of chickpea genome mapping, tagging of genes for ascochyta blight, fusarium wilt resistance and other traits, and requirements for MAS. Conventional breeding strategies to tolerate/avoid drought and chilling effects at flowering time, essential for changing from spring to winter sowing, are described. Recent approaches and future prospects for functional genomics of chickpea are discussed.     

Breeding drought tolerant cowpea: constraints,accomplishments, and future prospects

This review presents an overview of accomplishments on different aspects of cowpea breeding for drought tolerance. Furthermore it provides options to enhance the genetic potential of the crop by minimizing yield loss due to drought stress. Recent efforts have focused on the genetic dissection of drought tolerance through identification of markers defining quantitative trait loci (QTL) with effects on specific traits related to drought tolerance. Others have studied the relationship of the drought response and yield components, morphological traits and physiological parameters. To our knowledge, QTLs with effects on drought tolerance have not yet been identified in cowpea. The main reason is that very few researchers are working on drought tolerance in cowpea. Some other reasons might be related to the complex nature of the drought stress response, and partly to the difficulties associated with reliable and reproducible measurements of a single trait linked to specific molecular markers to be used for marker assisted breeding. Despite the fact that extensive research has been conducted on the screening aspects for drought tolerance in cowpea only very few—like the ‘wooden box’ technique—have been successfully used to select parental genotypes exhibiting different mechanisms of drought tolerance. Field and pot testing of these genotypes demonstrated a close correspondence between drought tolerance at seedling and reproductive stages. Some researchers selected a variety of candidate genes and used differential screening methods to identify cDNAs from genes that may underlie different drought tolerance pathways in cowpea. Reverse genetic analysis still needs to be done to confirm the functions of these genes in cowpea. Understanding the genetics of drought tolerance and identification of DNA markers linked to QTLs, with a clear path towards localizing chromosomal regions or candidate genes involved in drought tolerance will help cowpea breeders to develop improved varieties that combine drought tolerance with other desired traits using marker assisted selection.

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Evaluation of Yield Criteria for Drought and Heat Resistance in Chickpea (Cicer arietinum L.)

The chickpea (Cicer arietinum L.) is usually grown under rainfed, rather than irrigated conditions, where drought accompanied by heat stress is a major growth constraint. The aim of this study was to select chickpea genotypes having resistance to drought/heat stress and to identify the most appropriate selection criteria for this. A total of 377 chickpea accessions were sown 2 months later than normal for the Antalya region (Turkey) to increase their exposure to the drought and high‐temperature conditions of a typical summer in this part of the world. Interspersed between every 10 test genotypes as benchmark genotypes, were plants of the two known genotypes ILC 3279 (drought‐susceptible) and ILC 8617 (drought‐susceptible), while ICC 4958 (known drought‐resistant) and ICCV 96029 (known very early, double‐podded) were also sown for confirmation. All plants were subsequently screened for drought and heat stress resistance. Soon after the two known susceptible genotypes had died, evaluations of the entire trial were made visually on a scale from ‘1’ (free from drought/heat damage) to ‘9’ (all plants died from drought/heat). Yield loss in many of the test genotypes and in the two known susceptible genotypes (ILC 3279 and ILC 8617) rose to 100 %. The desi chickpeas (smaller, dark seeds) were generally more drought‐ and heat‐resistant than the kabuli chickpeas (larger, pale seeds). Two desi chickpeas, ACC 316 and ACC 317, were selected for drought and heat (>40 °C) resistance under field conditions. Seed weight was the trait least affected by adverse environmental conditions and having the highest heritability, and it should be used in early breeding selections. When breeding drought‐ and heat‐resistant chickpeas, path and multivariate analyses showed that days to the first flowering and maturity to escape terminal drought and heat stresses should be evaluated ahead of many other phenological traits, and harvest index, biological yield and pods per plant for increased yield should also be considered.     

Genotype selection for physiological responses of drought tolerance using molecular markers in polycross hybrids of orchardgrass

The present study aimed to assess the effect of contrasting levels of molecular and phenotypic diversity among polycross parents of orchardgrass on the performance of synthetic progeny with respect to physiological responses and drought tolerance. Four polycross groups each composed of six parental plants were evaluated under normal irrigation and drought stress conditions. A number of 923 inter simple sequence repeats and sequence related amplified polymorphism markers and several phenotypic traits were used to select contrasting levels of diversity (high and low) in parental genotypes. Highly significant correlation was observed between molecular distance and progeny performances at both normal irrigation and drought stress conditions. High molecular diversity among polycross parents led to a significant yield advantage of first generation progeny with averages of 34.40% for normal irrigation and 48.10% for drought stress conditions. Also crosses between genetically distant parents produced progeny with considerable drought tolerance and yield stability. Positive associations between phenotypic distance of parents and progeny performance were found for most physiological traits at both moisture regimes but phenotypic distances had weak association with forage yield, stress tolerance index and yield stability of progeny. Significant associations between drought tolerance index and some physiological traits confirmed the importance of these traits in conferring drought tolerance of orchardgrass. Our results underscore the effectiveness of marker‐assisted polycross breeding to improve drought tolerance and yield stability through physiological traits in orchardgrass.     

Independent and combined effects of high temperature and drought stress around anthesis on wheat

High temperature and drought stress are projected to reduce crop yields and threaten food security. While effects of heat and drought on crop growth and yield have been studied separately, little is known about the combined effect of these stressors. We studied detrimental effects of high temperature, drought stress and combined heat and drought stress around anthesis on yield and its components for three wheat cultivars originating from Germany and Iran. We found that effects of combined heat and drought on the studied physiological and yield traits were considerably stronger than those of the individual stress factors alone, but the magnitude of the effects varied for specific growth‐ and yield‐related traits. Single grain weight was reduced under drought stress by 13%–27% and under combined heat and drought stress by 43%–83% but not by heat stress alone. Heat stress significantly decreased grain number by 14%–28%, grain yield by 16%–25% and straw yield by 15%–25%. Cultivar responses were similar for heat but different for drought and combined heat and drought treatments. We conclude that heat stress as imposed in this study is less detrimental than the effects of those other studied stresses on growth and yield traits.     

Drought stress‐induced changes in starch yield and physiological traits in potato

Drought stress is an important limitation for potato (Solanum tuberosum L.) production as potato depends on appropriate water availability for high yields of good quality. Therefore, especially in the background of climate change, it is an important goal in potato breeding to improve drought stress tolerance. In this study, 34 European starch potato cultivars were evaluated for drought stress tolerance by growing under well‐watered and long‐term drought stress conditions in rainout shelters in 2 years’ pot trials. Besides yield, six physiological traits, that is free proline content, osmolality, total soluble sugar content, chlorophyll content (SPAD), cell membrane stability and crude protein content, were determined in leaves sampled during vegetative growth and during flowering to investigate their association with drought tolerance. ANOVA revealed significant treatment effects for all physiological traits and increased genotypic effects at flowering. The sensitivity of physiological traits to drought was significantly higher during flowering than during vegetative growth. Drought stress decreased starch yield significantly (p < .001), on average by 55%. Starch yield was significantly influenced by genotype and genotype × treatment interactions. Stress tolerance index (STI) calculated from starch yield ranged from 0.26 (sensitive) to 0.76 (tolerant) with significant genotype effects (p ≤ .001). STI correlated positively with cell membrane stability (r = .59) and crude protein content (r = .38) and negatively with osmolality (r = ?.57) and total soluble sugar content (r = ?.71). These contrary correlations suggest a dual adaptation strategy in potato under long‐term drought stress conditions including increased membrane stability combined with an increased osmolality due to an increased soluble sugar content.     

Development of early-flowering Kabuli chickpea with compound and simple leaves

Terminal drought is a major constraint to chickpea (Cicer arietinum L.) production. Autumn sowing and early flowering have been suggested as ways to benefit from the winter rains in short rainy seasons under dryland cropping. High‐yielding, late‐flowering, simple‐leafed (slv/slv) chickpea cultivars with good field resistance to Ascochyta blight have been bred recently. Changing plant architecture, by altering leaf shape, may affect agronomic performance. As no information is available on the effect of leaf shape on phenology and seed yield, this study was aimed at: (i) introducing the simple leaf trait into an early‐flowering chickpea background; (ii) comparing the grain yield of the two leaf types in early vs. late flowering backgrounds and (iii) producing breeding lines combining early flowering, large seeds and Ascochyta tolerance with both leaf types. Hybrid progeny were studied from the cross of ‘Sanford’ (slv/slv) and ICC7344, (compound, SLV/SLV). Four early‐podding, F₈ breeding lines were selected with either simple or compound leaves. In three different field experiments under dryland conditions (334–379 mm), they yielded ca. 1.4 t/ha as compared with 1.0 t/ha in the standard Israeli ‘Yarden’ on one site, but no significant differences in yield were obtained in the other two experiments.     

小麦种质资源抗旱性鉴定评价

培育抗旱节水小麦品种是保障我国粮食安全的重要途径之一, 优异抗旱种质资源筛选及抗旱性评价方法的研究对于提高抗旱育种效率具有关键作用。本研究采用反复干旱法和田间直接鉴定法分别鉴定323份小麦种质苗期和成株期的抗旱性。结果表明, 随着干旱次数的增加幼苗存活率逐渐下降, 而其变异系数和广义遗传力增加。成株期单株产量抗旱系数与综合抗旱性度量值D显著正相关(R ² = 0.609), 采用综合抗旱性度量值D有利于区分干旱对不同种质产量的影响力。苗期反复干旱存活率(DS)与单株产量的抗旱系数及综合抗旱性度量值D均无显著相关。基于反复干旱存活率筛选得到28份苗期强抗旱种质, 基于单株产量抗旱系数和综合抗旱性度量值D分别得到25和30份成株期强抗旱种质, 其中, 9份种质用2种评价方法均表现强抗旱; 21份种质在苗期和成株期均表现抗旱或强抗旱。本研究为小麦抗旱性评价方法及抗旱亲本的合理选择提供理论指导和信息支撑。     

Utilization of molecular markers for improving the phosphorus efficiency in crop plants

Phosphorus (P) is the second most growth limiting macronutrient after nitrogen and plays several important roles in all organisms including plants. In soil, P is available in both organic and inorganic forms. P deficiency reduces the growth and yield of several crop plants. Plants respond to P deficiency by the phenotypic changes especially by the modification of root architecture. Molecular marker‐assisted breeding (MAB) has been proposed as an important tool to identify and develop improved varieties of crop plants with efficient P‐use efficiency (PUE). Identification of quantitative trait loci (QTLs) for traits related to PUE has been considered as the first step in marker‐assisted selection (MAS) and improvement of crop yield programmes. In this review, we describe in detail on architectural changes of roots under P deficiency that are reported in various crops and discuss the efforts made to improve PUE using molecular marker tools. Details on QTLs identified for low P‐stress tolerance in various crop plants are presented. These QTLs can be used to improve PUE in crop plants through MAS and breeding, which may be beneficial to improve the yields under P‐deficient soil. Development of new and improved varieties using MAB will limit the use of non‐renewable fertilizers and improve PUE of key crop plants in low input agriculture.     

Recent progress in drought and salt tolerance studies in Brassica crops

Water deficit imposed by either drought or salinity brings about severe growth retardation and yield loss of crops. Since Brassica crops are important contributors to total oilseed production, it is urgently needed to develop tolerant cultivars to ensure yields under such adverse conditions. There are various physiochemical mechanisms for dealing with drought and salinity in plants at different developmental stages. Accordingly, different indicators of tolerance to drought or salinity at the germination, seedling, flowering and mature stages have been developed and used for germplasm screening and selection in breeding practices. Classical genetic and modern genomic approaches coupled with precise phenotyping have boosted the unravelling of genes and metabolic pathways conferring drought or salt tolerance in crops. QTL mapping of drought and salt tolerance has provided several dozen target QTLs in Brassica and the closely related Arabidopsis. Many drought- or salt-tolerant genes have also been isolated, some of which have been confirmed to have great potential for genetic improvement of plant tolerance. It has been suggested that molecular breeding approaches, such as marker-assisted selection and gene transformation, that will enhance oil product security under a changing climate be integrated in the development of drought- and salt-tolerant Brassica crops.     

Role of the maternal effect phenomena in improving water stress tolerance in narrow‐leafed lupine (Lupinus angustifolius)

This paper describes experiments concerning improvement of drought tolerance in narrow‐leafed lupine (Lupinus angustifolius L.) by maternal effects. The first step involved harvesting seeds from plants exposed and not exposed to drought during flowering and seed ripening. The next‐generation plants grown from these seeds were exposed to drought and the effects of this stress on their physiological processes were examined. To find out whether drought applied to parent plants may affect tolerance to this stress in progeny plants such features as plant growth, tissue water content, abscisic acid concentration and yield‐related parameters were assessed. The study revealed that the progeny plants grown from the seeds of drought‐treated plants were more tolerant to this stress than the plants grown from the seeds harvested from optimally watered maternal plants. Drought tolerance was manifested by a reduced concentration of abscisic acid, increased plant height and maintaining high leaf water content. Most importantly, these plants produced significantly higher yield when exposed to drought than the plants grown from the seeds harvested from optimally watered plants.     

Two Soybean Plant Introductions Display Slow Leaf Wilting and Reduced Yield Loss under Drought

Due to high costs of irrigation, limited availability of irrigation water in many locations and/or lack of irrigation capabilities, genetic improvement for drought tolerance is an effective method to reduce yield loss in soybean [Glycine max (L.) Merr.]. Slow wilting and minimal yield reduction under drought are important traits in evaluating drought tolerance. Two maturity group III soybean plant introductions (PIs, PI 567690 and PI 567731) and two elite cultivars (DKB38‐52 and Pana) were evaluated with and without irrigation on a sandy soil. Drought was imposed by withholding irrigation at full bloom and continued until moderate wilting was shown by the fast leaf wilting in the check cultivar, Pana. Then, irrigation was resumed until maturity. Genotypes were scored for leaf wilting during the stress period, and yields were assessed at the end of the growing season and used to calculate a drought index. Yields of the exotic PIs were lower than those of the checks under both drought and well‐watered conditions. However, the PIs exhibited significantly lower wilting and less yield loss under drought (higher drought index) than check cultivars. The two PIs may have useful genes to develop drought‐tolerant germplasm and cultivars and maybe useful in genetic and physiological studies to decipher mechanisms responsible for improving yield under limited water availability.     

Breeding banana (Musa spp.) for drought tolerance: A review

Drought is a major abiotic stress affecting banana production worldwide, leading to yield losses of up to 65%. Consequently, numerous efforts to understand and mitigate drought effects that include developing tolerant crop varieties are ongoing in several banana breeding programmes. The breeding efforts, however, have been greatly slowed down by inherent banana problems (polyploidy and male or female sterility) and complexity of drought tolerance (reportedly controlled by several genes). This review summarizes the pertinent research findings on water requirements of banana for its proper growth and productivity, symptoms of drought-sensitive varieties and field management strategies to cope with drought stress. The coping strategies deployed by resistant cultivars include high assimilation rates and water retention capacity as well as minor losses in leaf area and gaseous exchange. Reduced bunch weight, leaf chlorosis, wilting and strangled birth are underlined to be directly associated with drought susceptibility. Integration of conventional, molecular breeding and biotechnological tools as well as exploitation of the existing banana genetic diversity presents a huge opportunity for successful banana improvement.     

Aerial canopy temperature differences between fast‐ and slow‐wilting soya bean genotypes

Drought stress limits crop growth and yield in soya bean (Glycine max [L.] Merr.), but there are relatively few tools available to assess the ability of different genotypes to tolerate drought. Aerial infrared image analysis was evaluated as a potential tool for identifying drought tolerance in soya bean. Drought effects were evaluated from late vegetative to mid‐reproductive stages of soya bean development in an experiment with ten genotypes including five slow‐ and five fast‐wilting genotypes that were from a population derived from Benning×PI416937. There were two deficit irrigation levels for 2 years and one deficit irrigation level for the third year along with a fully irrigated control level. When the canopy was completely closed, relative canopy temperature was determined using an infrared camera taken from an aerial platform 50–75 m above the experiment. As water availability decreased, the relative canopy temperature generally increased. Moreover, slow‐wilting soya bean genotypes generally had lower canopy temperature compared to fast‐wilting genotypes, and grain yield was generally positively associated with cool canopy temperatures. The results indicate that the determination of canopy temperature is a promising tool for rapid characterization of drought‐related traits in soya bean.     

Assessing morphological characteristics of elite cotton lines from different breeding programmes for low temperature and drought tolerance

Cotton breeders in the United States strive to develop region‐specific genotypes adapted to low temperatures and variable soil moistures during early‐season planting. Nine elite upland cotton germplasm (Gossypium hirsutum L.) lines, representing public breeding programmes from nine states across the cotton belt, were evaluated for cold and drought stresses during seed germination and seedling growth stages. Lines were subjected to three treatments, such as low temperature well‐watered (22/14°C, WW), optimal temperature drought stress (30/22°C, DS) and optimal temperature well‐watered (30/22°C, WW; control), to examine genotypic variability for cold and drought tolerance. The treatment including drought stress was irrigated at 50% of the control. Shoot and root traits measured at 25 days after planting were significantly affected by drought and low temperature, where significant genetic variability among lines was observed for both shoot and root parameters. Response indices were developed to quantify variation in the degree of tolerance among the lines to low temperature and drought. Accordingly, OA‐33 was identified as the most low‐temperature‐tolerant line and Acala 1517‐99 as the most drought‐tolerant line. Identification of both cold‐ and drought‐tolerant genotypes suggests existing genotypic variability could provide breeders the opportunity to improve cultivar response to early‐season drought or cold conditions.     

Genetic Analysis of Grain Yield and Other Traits of Early‐Maturing Maize Inbreds under Drought and Well‐Watered Conditions

Maize (Zea mays L.) is an important staple food crop in West and Central Africa (WCA). However, its production is constrained by drought. Knowledge and understanding of the genetics of hybrid performance under drought is invaluable in designing breeding strategies for improving maize yield. One hundred and fifty hybrids obtained by crossing 30 inbreds in sets using the North Carolina Design II plus six checks were evaluated under drought and well‐watered conditions for 2 years at three locations in Nigeria. The objectives of the studies were to (i) determine the mode of gene action controlling grain yield and other important agronomic traits of selected early inbred lines, (ii) examine the relationship between per se performance of inbreds and their hybrids and (iii) identify appropriate testers for maize breeding programmes in WCA. General combining ability (GCA) and specific combining ability (SCA) mean squares were significant (P < 0.01) for grain yield and other traits under the research environments. The GCA accounted for 64.5 % and 62.3 % of the total variation for grain yield under drought and well‐watered conditions, indicating that additive gene action largely controlled the inheritance of grain yield of the hybrids. Narrow‐sense heritability was 67 % for grain yield under drought and 49 % under well‐watered conditions. The correlations between traits of early‐maturing parental lines and their hybrids were significant (P < 0.01) under drought, well‐watered and across environments. Mid‐parent and better‐parent heterosis for grain yield were 45.3 % and 18.4 % under drought stress and 111.9 % and 102.6 % under well‐watered conditions. Inbreds TZEI 31, TZEI 17, TZEI 129 and TZEI 157 were identified as the best testers. Drought‐tolerant hybrids with superior performance under stress and non‐stress conditions could be obtained through the accumulation of favourable alleles for drought tolerance in both parental lines.     

玉米耐旱性研究进展

干旱是目前影响玉米产量最重要的非生物胁迫因子。为了加快玉米耐旱性研究,提高育种效率,本研究归纳了近年来国内外玉米耐旱性研究现状将其总结为玉米耐旱性鉴定方法、遗传研究、品种选育以及耐旱性功能基因研究。对目前研究中存在的问题,如尚未有统一标准耐旱性鉴定体系、缺乏耐旱种质资源以及耐旱机理研究尚浅等进行了分析,并针对每个问题提出其解决对策,这为玉米耐旱性深入研究提供有力信息。     

White lupin (Lupinus albus) variation for adaptation to severe drought stress

This study aimed to reduce the gap of knowledge on white lupin drought tolerance variation, by assessing the grain yield of 21 landraces from major historical cropping regions, one variety and two breeding lines in a large phenotyping platform that imposed controlled severely stressed or moisture‐favourable conditions after an initial stage of favourable growth. Drought stress reduced grain yield by 79%. Genetic correlation coefficients indicated moderate consistency of genotype responses across conditions for grain yield (r_g = 0.76), fairly high consistency for straw yield (r_g = 0.85) and harvest index (r_g = 0.91), and high consistency for flowering time (r_g = 0.99). However, low genetic correlation for yield (r_g = 0.31) occurred among a subset of genotypes with early phenology. Specific adaptation to either condition implied significant (p = 0.05) genotype × condition interaction of crossover type between well‐performing genotypes. Early flowering was an important stress escape mechanism, but intrinsic drought tolerance could be inferred from responses of a few genotypes. Various landraces out‐yielded the improved germplasm under stressed or favourable conditions.