Strategies for Improving Drought Resistance in Grain Legumes

This review distills recent information on drought resistance characteristics of grain legumes with a view toward developing appropriate genetic enhancement strategies for water-limited environments. First, the possible adaptations that allow grain legumes to better cope with drought stress are summarized. It is suggested that there are considerable gains to be made in increasing yield and yield stability in environments characterized by terminal drought stress by further exploiting drought escape, by shortening crop duration. Many traits conferring dehydration avoidance and dehydration tolerance are available, but integrated traits, expressing at a higher level of organization, are suggested to be more useful in crop improvement programs. Possible genetic improvement strategies are outlined, ranging from empirical selection for yield in droughted environments to a physiological genetic approach. It is suggested that in view of recent advances in understanding drought resistance mechanisms, the latter strategy is becoming more feasible. It is concluded that use of this recently derived knowledge in a systematic manner can lead to significant gains in yield and yield stability of the world's major grain legumes, as they are mainly grown (and will continue to be grown) under rain-fed conditions.     

Efficiency of physiological trait-based and empirical selection approaches for drought tolerance in groundnut

Drought is the major abiotic constraint affecting groundnut productivity and quality worldwide. Most breeding programmes in groundnut follow an empirical approach to drought resistance breeding, largely based on kernel yield and traits of local adaptation, resulting in slow progress. Recent advances in the use of easily measurable surrogates for complex physiological traits associated with drought tolerance encouraged breeders to integrate these in their selection schemes. However, there has been no direct comparison of the relative efficiency of a physiological trait‐based selection approach (Tr) vis‐à‐vis an empirical approach (E) to ascertain the benefits of the former. The genetic material used in the present study originated from three common crosses and one institute‐specific cross from four collaborating institutes in India (total seven crosses). Each institute contributed six genotypes and each followed both the Tr and E selection approaches in each cross. The field trial of all selections, consisting of 192 genotypes (96 each Tr and E selections), was grown in 2000/2001 in a 4 × 48 alpha design in 12 season × location environments in India. The selection efficiency of Tr relative to E, RE_Tr, was estimated using the genetic concept of response to selection. Based on all the 12 environments, the two selection methods performed more or less similarly (RE_Tr= 1.045). When the 12 environments were grouped into rainy season and post‐rainy season, the relative response to selection in Tr method was higher in the rainy than in the post‐rainy season (RE_Tr= 1.220 vs 0.657) due to a higher genetic variance, lower G × E, and high h². When the 12 environments were classified into four clusters based on plant extractable soil‐water availability, the selection method Tr was superior to E in three of the four clusters (RE_Tr= 1.495, 0.612, 1.308, and 1.144) due to an increase in genetic variance and h² under Tr in clustered environments. Although the crosses exhibited significant differences for kernel yield, the two methods of selection did not interact significantly with crosses. Both methods contributed more or less equally to the 10 highest‐yielding selections (six for E and four for Tr). The six E selections had a higher kernel yield, higher transpiration (T), and nearly equal transpiration efficiency (TE) and harvest index (HI) relative to four Tr selections. The yield advantage in E selections came largely from greater T, which would likely not be an advantage in water‐deficient environments. From the results of these multi‐environment studies, it is evident that Tr method did not show a consistent superiority over E method of drought resistance breeding in producing a higher kernel yield in groundnut. Nonetheless, the integration of physiological traits (or their surrogates) in the selection scheme would be advantageous in selecting genotypes which are more efficient water utilisers or partitioners of photosynthates into economic yield. New biotechnological tools are being explored to increase efficiency of physiological trait‐based drought resistance breeding in groundnut.     

Models and tools for studying drought stress responses in peas

The pea (Pisum sativum L.) is an important pulse crop but the growing area is limited because of its relatively low yield stability. In many parts of the world the most important abiotic factor limiting the survival and yield of plants is the restricted water supply, and the crop productivity can only be increased by improving drought tolerance. Development of pea cultivars well adapted to dry conditions has been one of the major tasks in breeding programs. Conventional breeding of new cultivars for dry conditions required extensive selection and testing for yield performance over diverse environments using various biometrical approaches. Several morphological and biochemical traits have been proven to be related to drought resistance, and methods based on physiological attributes can also be used in development of better varieties. Osmoregulation plays a role in the maintenance of turgor pressure under water stress conditions, and information on the behaviour of genotypes under osmotic stress can help selection for drought resistance. Biotechnological approaches including in vitro test, genetic transformation, and the use of molecular markers and mutants could be useful tools in breeding of pea. In this minireview we summarized the present status of different approaches related to drought stress improvement in the pea.     

The relations between drought susceptibility index based on grain yield (DSIGY) and key physiological seedling traits in maize and triticale genotypes

The physiological reasons for the differences in sensitivity of C₃ and C₄ plant species to environmental stresses have not been thoroughly explained. In this study the effects of drought stress on the growth and selected physiological traits were examined in the seedlings of 13 single cross maize (C₄ plant) hybrids and 11 spring triticale (C₃ plant) breeding lines and varieties differing in drought sensitivity. For plants in the seedling stage the results demonstrated a genetic variation in dry matter accumulation of shoots and roots (DW_S, DW_R), number (N) and length (L) of particular components (seminal, seminal adventitious, nodal) of the root system, membrane injury by soil drought (LI_D), osmotic and high temperature stress (LI_OS, LI_HT), water potential (ψ), water loss (WL), grain germination in osmotic stress (FG, PI), and seedling survival (SS). Seedlings grown under moderate soil drought showed a decrease in dry matter of the top parts and roots and a decrease in the length of seminal, seminal adventitious and nodal roots in comparison to seedlings grown in control conditions. The observed harmful effects of drought stress were more distinct in drought sensitive genotypes. Used in this paper drought susceptibility indexes (DSI_GY) were calculated in other experiment by determining the changes in grain yield (GY) under two soil moisture levels (irrigated and drought). The variation of DSI_GY for maize ranges from 0.381 to 0.650 and for triticale from 0.354 to 0.578. The correlations between DSI_GY and laboratory tests (LI, FG, SS) confirmed that they are good indicators of drought tolerance in plants. The highest values of genetic variation were observed in LI, DW_S, SS and WL and the lowest in the measurements of ψ FG, PI, L_S, L_SA and L_N. The correlation coefficients between LI_OS and LI_HT tests were, in most of the considered cases, statistically significant, which indicates that in maize and triticale the mechanisms of membrane injury caused by simulated drought or high temperature are physiologically similar. It can be concluded that an approach to the breeding of maize and triticale for drought tolerance using these tests can be implemented on the basis of separate selection for each trait or for all of them simultaneously. In that case, it would be necessary to determine the importance of the trait in relation to growth phase, drought timing and level, as well as its associations with morphological traits contributing to drought tolerance. The obtained values of the correlation coefficient between laboratory tests suggest that the same physiological traits may be applied as selection criteria in drought tolerance of maize and triticale genotypes.     

Drought Stress in Wheat during Flowering and Grain-filling Periods

Drought is a major environmental stress threatening wheat productivity worldwide. Global climate models predict changed precipitation patterns with frequent episodes of drought. Although drought impedes wheat performance at all growth stages, it is more critical during the flowering and grain-filling phases (terminal drought) and results in substantial yield losses. The severity and duration of the stress determine the extent of the yield loss. The principal reasons for these losses are reduced rates of net photosynthesis owing to metabolic limitations—oxidative damage to chloroplasts and stomatal closure—and poor grain set and development. A comprehensive understanding of the impact of terminal drought is critical for improving drought resistance in wheat, with marker-assisted selection being increasingly employed in breeding for this resistance. The limited success of molecular breeding and physiological strategies suggests a more holistic approach, including interaction of drought with other stresses and plant morphology. Furthermore, integration of physiological traits, genetic and genomic tools, and transgenic approaches may also help to improve resistance against drought in wheat. In this review, we describe the influence of terminal drought on leaf senescence, carbon fixation, grain set and development, and explain drought resistance mechanisms. In addition, recent developments in integrated approaches such as breeding, genetics, genomics, and agronomic strategies for improving resistance against terminal drought in wheat are discussed.     

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.     

The development and application of molecular markers for abiotic stress tolerance in barley

This article represents some current thinking and objectives in the use of molecular markers to abiotic stress tolerance. Barley has been chosen for study as it is an important crop species, as well as a model for genetic and physiological studies. It is an important crop and, because of its well-studied genetics and physiology, is an excellent candidate in which to devise more efficient breeding methods. Abiotic stress work on cultivated gene pools of small grain cereals frequently shows that adaptive and developmental genes are strongly associated with responses. Developmental genes have strong pleiotropic effects on a number of performance traits, not just abiotic stresses. One concern is that much of the genetic variation for improving abiotic stress tolerance has been lost during domestication, selection and modern breeding, leaving pleiotropic effects of the selected genes for development and adaptation. Such genes are critical in matching cultivars to their target agronomic environment, and since there is little leverage in changing these, other sources of variation may be required. In barley, and many other crops, greater variation to abiotic stresses exists in primitive landraces and related wild species gene pools. Wild barley, Hordeum spontaneum C. Koch is the progenitor of cultivated barley, Hordeum vulgare L. and is easily hybridized to H. vulgare. Genetic fingerprinting of H. spontaneum has revealed genetic marker associations with site-of-origin ecogeographic factors and also experimentally imposed stresses. Genotypes and collection sites have been identified which show the desired variation for particular stresses. Doubled haploid and other segregating populations, including landrace derivatives have been used to map genetically the loci involved. These data can be used in molecular breeding approaches to improve the drought tolerance of barley. One strategy involves screening for genetic markers and physiological traits for drought tolerance, and the associated problem of drought relief-induced mildew susceptibility in naturally droughted fields of North Africa.     

The evaluation of photosynthetic parameters in maize inbred lines subjected to water deficiency: Can these parameters be used for the prediction of performance of hybrid progeny?

The response of selected photosynthetic and morphological parameters of plants to drought was examined in 5 inbred lines of maize (Zea mays L.) and their 10 F1 hybrids. The aim of the study was to establish whether the photosynthetic performance of parental genotypes under drought conditions correlates with the performance of their progeny and whether the net photosynthetic rate, the chlorophyll fluorescence parameters or the content of photosynthetic pigments could be used as reliable physiological markers for early breeding generations. The relative importance of the additive and the nonadditive (dominance, maternal) genetic effects in the inheritance of these parameters was also assessed by means of the quantitative genetics analysis. The results showed that the nonadditive genetic effects associated with a particular combination of genotypes or a particular direction of crossing are at least equally and often even more important as the additivity and that these genetic effects almost totally change with the exposure of plants to drought conditions. This was reflected in the inability to predict the response of F1 hybrids to drought on the basis of the photosynthetic performance of their parents, which indicates that the practical usability of such parameters in maize breeding programs is rather limited.     

鱼和熊掌的选择:反向重复序列变异介导的玉米环境适应与产量平衡

作物育种的目标是找到产量和抗性的最佳平衡点,其中涉及“鱼和熊掌”二者兼得的选择策略。哪些逆境负调控位点影响产量性状,以及如何调控等是突破育种瓶颈的重要科学问题。近百年来,高产玉米(Zeamays)育种使玉米单产不断提高,同时现代玉米品种对干旱的敏感性也呈现出增强趋势,故而存在高产稳产的潜在风险。可对于这一现象背后确切的遗传机制却知之甚少,从而限制了既高产又高抗玉米新品种的培育。玉米的非生物胁迫抗性与产量性状均为多基因控制的复杂数量性状,涉及全基因组范围内大量基因的表达与调控。玉米基因组内存在大量的小RNA (sRNA),其对基因表达起精细调控作用,但人们对sRNA调控作物环境胁迫应答与产量性状机制的理解仍然有限。近日,华中农业大学代明球课题组与李林和李峰两个课题组合作,基于对338份玉米关联群体在不同环境下的sRNA表达组分析,鉴定到大量干旱应答的sRNA,以及调控这些sRNA表达的遗传位点(eQTL);并克隆了8号染色体上1个干旱特异性eQTL热点DRESH8。生物信息学分析显示, DRESH8是1个由转座子组成的长度约为21.4 kb的反向重复序列(TE-IR)。DRESH8通过产...     

Genotype and phenotype associations with drought tolerance in barley tested in North Africa

A review is presented of genetic strategies deployed in a 3-yr project on drought tolerance in barley. Data were collected on genetic, physiological and agronomic traits in non-irrigated and irrigated field trials in Egypt, Morocco and Tunisia. A wide range of barley germplasm (developed from African and European cultivars, adapted landraces and wild barleys) was tested, and positive traits were found in each gene pool. The contrasting environments of the three North African countries had major effects on plant/genotype performance. Genetic effects were also detected, as were genotype × environment interactions. A range of strategies were deployed to investigate the physiology and genetics of quantitative traits associated with field performance. Quantitative trait locus (QTL) analysis was performed using backcross lines, recombinant inbred lines and doubled haploid mapping populations. A detailed genetic map was generated in the Tadmor × (ER/Apm) recombinant inbred lines, an important mapping population specifically developed by ICARDA (Centre for Agricultural Research in Dry Areas) and CIMMYT (International Maize and Wheat Improvement Center) to study drought. Quantitative trait loci (QTLs) for grain yield and other important morphological and physiological traits were also identified in a population of doubled haploids derived from F2BCj plants from a cross between a cultivar and a wild barley accession. Significantly, the wild parental line was found to contribute a number of positive alleles for yield. Effects of major developmental genes could explain many of the responses observed. QTLs were found to cluster around major genes controlling flowering time (sghI), plant stature (sdwI and arie.GP) and ear type (vrsl), and it is highly likely that the associations represent pleiotropic effects. Some QTLs were associated with candidate genes such as dehydrins and rubisco activase. One of the most significant results was the identification and generation of material that out performed the best local standards in the three participating North African countries; the selected lines have now entered local breeding programmes. The strategies adopted provided information on physiological traits, genotypes and genetic markers that could be used for marker-assisted selection. Target QTLs and their associated genetic markers may be deployed in marker assisted selection programmes to match crop phenology to the field environment.     

Genomics-based approaches to improve drought tolerance of crops

The genetic bases of the molecular, cellular and developmental responses to drought involve many gene functions regulated by water availability. Genomics-based approaches provide access to agronomically desirable alleles present at quantitative trait loci (QTLs) that affect such responses, thus enabling us to improve the drought tolerance and yield of crops under water-limited conditions more effectively. Marker-assisted selection is already helping breeders improve drought-related traits. Analysis of sequence data and gene products should facilitate the identification and cloning of genes at target QTLs. Based on such premises, we envision a quick broadening of our understanding of the genetic and functional basis of drought tolerance. Novel opportunities will be generated for tailoring new genotypes "by design". Harnessing the full potential of genomics-assisted breeding will require a multidisciplinary approach and an integrated knowledge of the molecular and physiological processes influencing tolerance to drought.     

Translational research impacting on crop productivity in drought-prone environments

Conventional breeding for drought-prone environments (DPE) has been complemented by using exotic germplasm to extend crop gene pools and physiological approaches that consider water uptake (WU), water-use efficiency (WUE), and harvest index (HI) as drivers of yield. Drivers are associated with proxy genetic markers, such as carbon-isotope discrimination for WUE, canopy temperature for WU, and anthesis-silking interval for HI in maize. Molecular markers associated with relevant quantitative trait loci are being developed. WUE has also been increased through combining understanding of root-to-shoot signaling with deficit irrigation. Impacts in DPE will be accelerated by combining proven technologies with promising new strategies such as marker-assisted selection, and genetic transformation, as well as conservation agriculture that can increase WU while averting soil degradation.     

Quantitative Trait Loci Mapping of Maize Yield and Its Components Under Different Water Treatments at Flowering Time

Drought or water stress is a serious agronomic problem resulting in maize (Zea mays L.) yield loss throughout the world. Breeding hybrids with drought tolerance is one important approach for solving this problem. However, lower efficiency and a longer period of breeding hybrids are disadvantages of traditional breeding programs. It is generally recognized that applying molecular marker techniques to traditional breeding programs could improve the efficiency of the breeding of drought‐tolerant maize. To provide useful information for use in studies of maize drought tolerance, the mapping and tagging of quantitative trait loci (QTL) for yield and its components were performed in the present study on the basis of the principle of a mixed linear model. Two hundred and twenty‐one recombinant inbred lines (RIL) of Yuyu 22 were grown under both well‐watered and water‐stressed conditions. In the former treatment group, plants were well irrigated, whereas those in the latter treatment group were stressed at flowering time. Ten plants of each genotype were grown in a row that was 3.00 m × 0.67 m (length × width). The results show that a few of the QTL were the same (one additive QTL for ear length, two additive QTL and one pair of epistatic QTL for kernel number per row, one additive QTL for kernel weight per plant), whereas most of other QTL were different between the two different water treatment groups. It may be that genetic expression differs under the two different water conditions. Furthermore, differences in the additive and epistatic QTL among the traits under water‐stressed conditions indicate that genetic expression also differs from trait to trait. Major and minor QTL were detected for the traits, except for kernel number per row, underwater‐stressed conditions. Thus, the genetic mechanism of drought tolerance in maize is complex because the additive and epistatic QTL exist at the same time and the major and minor QTL all contribute to phenotype under water‐stressed conditions. In particular, epidemic QTL under water‐stressed conditions suggest that it is important to investigate the drought tolerance of maize from a genetic viewpoint. (Managing editor: Wei Wang)     

Genomic models with genotype × environment interaction for predicting hybrid performance: an application in maize hybrids

Key message

A new genomic model that incorporates genotype?×?environment interaction gave increased prediction accuracy of untested hybrid response for traits such as percent starch content, percent dry matter content and silage yield of maize hybrids.

Abstract

The prediction of hybrid performance (HP) is very important in agricultural breeding programs. In plant breeding, multi-environment trials play an important role in the selection of important traits, such as stability across environments, grain yield and pest resistance. Environmental conditions modulate gene expression causing genotype?×?environment interaction (G?×?E), such that the estimated genetic correlations of the performance of individual lines across environments summarize the joint action of genes and environmental conditions. This article proposes a genomic statistical model that incorporates G?×?E for general and specific combining ability for predicting the performance of hybrids in environments. The proposed model can also be applied to any other hybrid species with distinct parental pools. In this study, we evaluated the predictive ability of two HP prediction models using a cross-validation approach applied in extensive maize hybrid data, comprising 2724 hybrids derived from 507 dent lines and 24 flint lines, which were evaluated for three traits in 58 environments over 12 years; analyses were performed for each year. On average, genomic models that include the interaction of general and specific combining ability with environments have greater predictive ability than genomic models without interaction with environments (ranging from 12 to 22%, depending on the trait). We concluded that including G?×?E in the prediction of untested maize hybrids increases the accuracy of genomic models.

     

Root and leaf traits, water use and drought tolerance of maize genotypes

The main components of drought tolerance of six maize genotypes were studied to evaluate crop performance in water limiting environments: (1) the postponement of dehydration by reduced transpiration rate (TR) and an increased efficiency of water acquisition from soil; (2) the tolerance of dehydration by effective physiological water use. The aim was to describe the genotype dependent response to drought in leaf and root traits and water relations using data from controlled environment and field experiments, and using dynamic simulation by the Swedish Coup model. High genetic variation was detected in the root density, acquisition efficiency and water use among the genotypes. The female parent lines had the greatest TR with the smallest dry matter accumulation in water deficiency, whereas hybrids could acquire more water from dryer soil while maintaining a lower TR. Hybrid Mv 444 increased water potential more strongly in leaves than hybrid Norma. The postponement of dehydration was observed for Norma, while more tolerance to dehydration characterized Mv 444. Simulation was an effective tool for testing hypotheses considering water acquisition efficiency and for summarizing the results of the measurements in a formalized structure; it helped to quantify the dynamics of water availability and the impact of drought on the growth of the maize genotypes.     

Quantitative trait loci influencing drought tolerance in grain sorghum (Sorghum bicolor L. Moench)

Drought is a major constraint in sorghum production worldwide. Drought-stress in sorghum has been characterized at both pre-flowering and post-flowering stages resulting in a drastic reduction in grain yield. In the case of post-flowering drought stress, lodging further aggravates the problem resulting in total loss of crop yield in mechanized agriculture. The present study was conducted to identify quantitative trait loci (QTLs) controlling post-flowering drought tolerance (stay green), pre-flowering drought tolerance and lodging tolerance in sorghum using an F₇ recombinant inbred line (RIL) population derived from the cross SC56×Tx7000. The RIL lines, along with parents, were evaluated for the above traits in multiple environments. With the help of a restriction fragment length polymorphism (RFLP) map, which spans 1,355 cM and consists of 144 loci, nine QTLs, located over seven linkage groups were detected for stay green in several environments using the method of composite interval mapping. Comparison of the QTL locations with the published results indicated that three QTLs located on linkage groups A, G and J were consistent. This is considered significant since the stay green line SC56 used in our investigation is from a different source compared to B35 that was used in all the earlier investigations. Comparative mapping has shown that two stay green QTLs identified in this study corresponded to stay green QTL regions in maize. These genomic regions were also reported to be congruent with other drought-related agronomic and physiological traits in maize and rice, suggesting that these syntenic regions might be hosting a cluster of genes with pleiotropic effects implicated in several drought tolerance mechanisms in these grass species. In addition, three and four major QTLs responsible for lodging tolerance and pre-flowering drought tolerance, respectively, were detected. This investigation clearly revealed the important and consistent stay green QTLs in a different stay green source that can logically be targeted for positional cloning. The identification of QTLs and markers for pre-flowering drought tolerance and lodging tolerance will help plant breeders in manipulating and pyramiding those traits along with stay green to improve drought tolerance in sorghum. Received: 2 June 2000 / Accepted: 15 November 2000     

Physiological responses of groundnut (Arachis hypogea L.) to drought stress and its amelioration: a critical review

Groundnut (Arachis hypogea L.), is an important legume cash crop for the tropical farmers and its seeds contain high amounts of edible oil (43–55%) and protein (25–28%). Even though it is a fairly drought-tolerant, production fluctuates considerably as a result of rainfall variability. To develop a water stress response function in groundnut, research works have been done to improve the performance under varying degrees of stress at various physiological stages of crop growth. This review summarizes recent information on drought resistance characteristics of groundnut with a view toward developing appropriate genetic enhancement strategies for water-limited environments. It is suggested that there are considerable gains to be made in increasing yield and stabilizing the yield in environments characterized by terminal drought stress and by shortening crop duration. Many traits conferring dehydration avoidance and dehydration tolerance are available, but integrated traits, expressing at a high level of organization are suggested to be more useful in crop improvement programs. Possible genetic improvement strategies are outlined, ranging from empirical selection for yield in drought environments to a physiological–genetic approach. It was also suggested that in view of recent advances in understanding drought resistance mechanisms, the later strategy is becoming more feasible. It is summarized that application of knowledge into practice in a systematic manner can lead to significant gains in yield and yield stability of the worlds groundnuts production. Research is needed to develop transferable technology to help farmers of arid and semi-arid regions. Increasing soil moisture storage by soil profile management and nutrient management for quick recovery from drought are some of the areas that need to be explored further.