Challenges in pea breeding for tolerance to drought: Status and prospects

Drought is increasingly frequent in the context of climate change and is considered a major constraint for crop yield. Water scarcity can impair growth, disturb plant water relations and reduce water use efficiency. Pea (Pisum sativum) is a temperate grain legume rich in protein, fibre, micronutrients and bioactive compounds that can benefit human health. In reducing pea yield because of drought, the intensity and duration of stress are critical. This review describes several drought resistance mechanisms in pea based on morphology, physiology and biochemical changes during/after the water deficit period. Drought tolerance of pea can be managed by adopting strategies such as screening, breeding and marker-assisted selection. Therefore, various biotechnological approaches have led to the development of drought-tolerant pea cultivars. Finally, the main objective of the current research is to point out some useful traits for drought tolerance in peas and also, mention the methods that can be useful for future studies and breeding programmes.     

Label‐free quantitative proteomic analysis of tolerance to drought in Pisum sativum

Abiotic stresses caused by adverse environmental conditions are responsible for heavy economic losses on pea crop, being drought one of the most important abiotic constraints. Development of pea cultivars well adapted to dry conditions has been one of the major tasks in breeding programs. The increasing food requirements drive the necessity to broaden the molecular basis of tolerance to drought to develop pea cultivars well adapted to dry conditions. We have used a shotgun proteomic approach (nLC‐MSMS) to study the tolerance to drought in three pea genotypes that were selected based on differences in the level of water deficit tolerance. Multivariate statistical analysis of data unraveled 367 significant differences of 700 identified when genotypes and/or treatment were compared. More than half of the significantly changed proteins belong to primary metabolism and protein regulation categories. We propose different mechanisms to cope drought in the genotypes studied. Maintenance of the primary metabolism and protein protection seems a strategy for drought tolerance. On the other hand susceptibility might be related to maintenance of the homeostatic equilibrium, a very energy consuming process. Data are available via ProteomeXchange with identifier PXD004587.     

抗旱相关基因在烟草中的应用研究进展

干旱是影响烟草正常生长、发育、产量和烟叶品质的一个重要逆境因子。在干旱胁迫下,植物体内会通过激发一些抗旱基因的表达来增强植物的抗旱能力。目前,很多抗旱相关的功能蛋白基因和调控蛋白基因已被克隆并在烟草中实现了遗传转化,外源抗旱基因的表达提高了转基因烟草的抗旱能力。抗旱基因的克隆为烟草抗旱新品种的培育奠定了良好的分子基础,系统深入地研究抗旱相关基因在干旱胁迫条件下的表达与调控,可为通过基因工程手段提高烟草的抗旱能力开辟新途径,同时也能为其他农作物的抗旱分子育种和品种改良提供基因资源。     

Advances in Drought Resistance of Rice

Water deficit is a serious environmental stress and the major constraint to rice productivity. Losses in rice yield due to water shortage probably exceed losses from all other causes combined and the extent of the yield loss depends on both the severity and duration of the water stress. Drought affects rice at morphological, physiological, and molecular levels such as delayed flowering, reduced dry matter accumulation and partitioning, and decreased photosynthetic capacity as a result of stomatal closure, metabolic limitations, and oxidative damage to chloroplasts. Small-statured rice plants with reduced leaf area and short growth duration are better able to tolerate drought stress, although the mechanisms are not yet fully understood. Increased water uptake by developing larger and deeper root systems, and the accumulation of osmolytes and osmoprotectants are other important mechanisms for drought resistance. Drought resistance in rice has been improved by using plant growth regulators and osmoprotectants. In addition, several enzymes have been found that act as antioxidants. Silicon has also improved drought resistance in rice by silicification of the root endodermis and improving water uptake. Seed priming improves germination and crop stand establishment under drought. Rice plants expressing HVA1, LEA proteins, MAP kinase, DREB and endo-1, 3-glucanase are better able to withstand drought stress. Polyamines and several enzymes act as antioxidants and reduce adverse effects of drought stress in rice. Drought resistance can be managed by developing and selecting drought-tolerant genotypes. Rice breeding and screening may be based on growth duration, root system, photosynthesis traits, stomatal frequency, specific leaf weight, leaf water potential, and yield in target environments. This review discusses recent developments in integrated approaches, such as genetics, breeding and resource management to increase rice yield and reduce water demand for rice production.     

Inducing drought tolerance in plants: Recent advances

Undoubtedly, drought is one of the prime abiotic stresses in the world. Crop yield losses due to drought stress are considerable. Although a variety of approaches have been used to alleviate the problem of drought, plant breeding, either conventional breeding or genetic engineering, seems to be an efficient and economic means of tailoring crops to enable them to grow successfully in drought-prone environments. During the last century, although plant breeders have made ample progress through conventional breeding in developing drought tolerant lines/cultivars of some selected crops, the approach is, in fact, highly time-consuming and labor- and cost-intensive. Alternatively, marker-assisted breeding (MAB) is a more efficient approach, which identifies the usefulness of thousands of genomic regions of a crop under stress conditions, which was, in reality, previously not possible. Quantitative trait loci (QTL) for drought tolerance have been identified for a variety of traits in different crops. With the development of comprehensive molecular linkage maps, marker-assisted selection procedures have led to pyramiding desirable traits to achieve improvements in crop drought tolerance. However, the accuracy and preciseness in QTL identification are problematic. Furthermore, significant genetic × environment interaction, large number of genes encoding yield, and use of wrong mapping populations, have all harmed programs involved in mapping of QTL for high growth and yield under water limited conditions. Under such circumstances, a transgenic approach to the problem seems more convincing and practicable, and it is being pursued vigorously to improve qualitative and quantitative traits including tolerance to biotic and abiotic stresses in different crops. Rapid advance in knowledge on genomics and proteomics will certainly be beneficial to fine-tune the molecular breeding and transformation approaches so as to achieve a significant progress in crop improvement in future. Knowledge of gene regulation and signal transduction to generate drought tolerant crop cultivars/lines has been discussed in the present review. In addition, the advantages and disadvantages as well as future prospects of each breeding approach have also been discussed.     

Defining selection criteria to improve yield under drought

The many selection criteria that have been proposed to increase drought resistance of our crops have had little, if any, impact on improving crop yields in dry environments. There are several likely reasons for this lack of success. Some of these are: (i) criteria proposed have been related more to survival mechanisms under drought than to productivity, (ii) criteria are inappropriate to the target environment, and (iii) criteria are temporal and are therefore likely to have minimal impact on growth and yield over the entire lifecycle. Another important reason is that breeders have not been convinced the proposed criteria will be successful as they are too difficult to measure. On the other hand, empirical breeding programmes to improve yield under drought have been successful. Surprisingly, some of the greatest successes have been achieved by breeding in environments where water is non-limiting. This paper reviews breeding approaches to improve yield under drought. It focuses on critical factors that must be considered to identify likely plant attributes that can be targeted. These factors, their link with yield, the nature of the target environment, the level of organisation where the trait is expressed are discussed. Three quite different examples are given to emphasize the above considerations and which show substantial promise in targeting traits to improve yield under drought. They are drought at flowering, improving transpiration efficiency and improving early leaf area development.     

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(Zea mays)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通过产生小干扰RNA (siRNA)介导抗旱基因的转录后沉默, 并间接抑制产量负调控因子的表达, 在负调控干旱应答的同时正调控产量性状。进一步研究发现, DRESH8在玉米驯化和改良过程中受到了人工选择。据此, 他们认为DRESH8可能是玉米平衡抗旱性和产量的关键遗传位点。该研究在全基因组水平上揭示了作物调控产量和环境胁迫抗性平衡的关键遗传机制, 同时也鉴定到大量IR位点, 为未来“高抗、高产”玉米设计育种提供了有价值的操控靶点。     

Seasonal radiation interception, dry matter production and yield determination for a semi-leafless pea (Pisum sativum) breeding selection under contrasting field conditions

Radiation interception, dry matter accumulation, flower and pod production and yield were measured for a semi-leafless pea (Pisum sativum) breeding selection (BS3) on three contrasting sites. Differences in soil moisture availability were largely responsible for a three-fold difference in yield between sites. Radiation interception was related to dry matter production by calculating photosynthetic efficiencies. In the absence of lodging, crop canopies converted intercepted radiation into dry matter with constant efficiency (?) throughout the season; under conditions of moisture stress ? was reduced. Serious lodging during the post-flowering period on one site resulted in a mean seasonal photosynthetic efficiency (?) 17% lower than ?. The ability of the pea crop canopy to intercept radiation was related also to yield components.     

Interaction of aluminium and drought stress on root growth and crop yield on acid soils

Background

Aluminium (Al) toxicity and drought stress are two major constraints for crop production in the world, particularly in the tropics. The variation in rainfall distribution and longer dry spells in much of the tropics during the main growing period of crops are becoming increasingly important yield-limiting factors with the global climate change. As a result, crop genotypes that are tolerant of both drought and Al toxicity need to be developed.

Scope

The present review mainly focuses on the interaction of Al and drought on root development, crop growth and yield on acid soils. It summarizes evidence from our own studies and other published/related work, and provides novel insights into the breeding for the adaptation to these combined abiotic stresses. The primary symptom of Al phytotoxicity is the inhibition of root growth. The impeded root system will restrict the roots for exploring the acid subsoil to absorb water and nutrients which is particularly important under condition of low soil moisture in the surface soil under drought. Whereas drought primarily affects shoot growth, effects of phytotoxic Al on shoot growth are mostly secondary effects that are induced by Al affecting root growth and function, while under drought stress root growth may even be promoted. Much progress has recently been made in the understanding of the physiology and molecular biology of the interaction between Al toxicity and drought stress in common bean (Phaseolus vulgaris L.) in hydroponics and in an Al-toxic soil.

Conclusions

Crops growing on acid soils yield less than their potential because of the poorly developed root system that limits nutrient and water uptake. Breeding for drought resistance must be combined with Al resistance, to assure that drought resistance is expressed adequately in crops grown on soils with acid Al-toxic subsoils.     

Assessment of yield based selection under managed field stress condition for breeding for rice yield improvement under drought

The progress in development and dissemination of drought tolerant lines has been slow as compared to the increasing drought prevalence in the rice growing regions. Significant amount of work has been done in the past on drought resistance traits in rice crop, still the benefit of improved drought tolerant rice cultivars reaching the farmer’s field is not very high and ways to expedite the development of drought tolerant and productive rice cultivars needs to be addressed. In this article, an assessment of easily practicable approach of managed stress screening and prospect of direct selection for yield under drought stress is discussed. Also the large effect yield QTLs identified for grain yield under drought stress field conditions is being reviewed for successful introgression into elite genetic background for developing drought tolerant cultivars with improved yield for the drought prone target environment.     

Characterizing drought stress and trait influence on maize yield under current and future conditions

Global climate change is predicted to increase temperatures, alter geographical patterns of rainfall and increase the frequency of extreme climatic events. Such changes are likely to alter the timing and magnitude of drought stresses experienced by crops. This study used new developments in the classification of crop water stress to first characterize the typology and frequency of drought‐stress patterns experienced by European maize crops and their associated distributions of grain yield, and second determine the influence of the breeding traits anthesis‐silking synchrony, maturity and kernel number on yield in different drought‐stress scenarios, under current and future climates. Under historical conditions, a low‐stress scenario occurred most frequently (ca. 40%), and three other stress types exposing crops to late‐season stresses each occurred in ca. 20% of cases. A key revelation shown was that the four patterns will also be the most dominant stress patterns under 2050 conditions. Future frequencies of low drought stress were reduced by ca. 15%, and those of severe water deficit during grain filling increased from 18% to 25%. Despite this, effects of elevated CO₂ on crop growth moderated detrimental effects of climate change on yield. Increasing anthesis‐silking synchrony had the greatest effect on yield in low drought‐stress seasonal patterns, whereas earlier maturity had the greatest effect in crops exposed to severe early‐terminal drought stress. Segregating drought‐stress patterns into key groups allowed greater insight into the effects of trait perturbation on crop yield under different weather conditions. We demonstrate that for crops exposed to the same drought‐stress pattern, trait perturbation under current climates will have a similar impact on yield as that expected in future, even though the frequencies of severe drought stress will increase in future. These results have important ramifications for breeding of maize and have implications for studies examining genetic and physiological crop responses to environmental stresses.     

The impact of different soil moisture and soil compaction on the growth of triticale root system

Effects of different soil moisture (soil drought and waterlogging) and soil compaction (1.33 and 1.50 g·cm⁻³) on the growth and morphological traits of the root system were studied in four breeding forms and seven cultivars of triticale. Morphological changes, including the restriction of root extension, expansion and proliferation of laterals roots, occur in plants grown in different soil moisture and in compact soil. The investigations comprised quantitative and qualitative analyses of a developed plant root system through determining the number, length and dry matter of the particular components of the root system. Obtained results have demonstrated a relatively broad variation in the habit of the triticale root system. Plants grown under compact soil and low or high soil water content showed a smaller number and less dry matter of lateral branching than plants grown in control conditions. The harmful effects of compact soil and drought conditions on the growth of roots was greater when compared with that of plants exposed to waterlogging. The observed effects of all treatments were more distinct in a drought sensitive strains. The drought resistant forms were a more characterize with extensive rooting and by smaller alterations in the root morphology under the stress conditions compared with drought sensitive one. Results confirm that the breeding forms (CHD-12 and CHD-173) of a high drought susceptibility was found to be also more sensitive to periodical soil water excess. A more efficient water use and a lower shoot to root (S/R) ratio were found to be major reasons for a higher stress resistance of the breeding forms (CHD-220 and CHD-247). The reasons for a different response of the examined breeding forms and cultivars to the conditions of drought or waterlogging may be a more economical water balance and more favourable relations between the shoot and root dimensions in the drought resistant forms and cultivars. The results suggest that the morphological traits of the triticale root system may be used in practice as direct or indirect selection criteria in maize breeding.     

Plant Breeding and Drought in C3 Cereals: What Should We Breed For?

Drought is the main abiotic constraint on cereal yield. Analysingphysiological determinants of yield responses to water may helpin breeding for higher yield and stability under drought conditions.The traits to select (either for stress escape, avoidance ortolerance) and the framework where breeding for drought stressis addressed will depend on the level and timing of stress inthe targeted area. If the stress is severe, breeding under stress-freeconditions may be unsuccessful and traits that confer survivalmay become a priority. However, selecting for yield itself understress-alleviated conditions appears to produce superior cultivars,not only for optimum environments, but also for those characterizedby frequent mild and moderate stress conditions. This impliesthat broad avoidance/tolerance to mild–moderate stressesis given by constitutive traits also expressed under stress-freeconditions. In this paper, we focus on physiological traitsthat contribute to improved productivity under mild–moderatedrought. Increased crop performance may be achieved throughimprovements in water use, water-use efficiency and harvestindex. The first factor is relevant when soil water remainsavailable at maturity or when deep-rooted genotypes access waterin the soil profile that is not normally available; the twolatter conditions become more important when all available wateris exhausted by the end of the crop cycle. Independent of themechanism operating, a canopy able to use more water than anotherwould have more open stomata and therefore higher canopy temperaturedepression, and ¹³C discrimination (¹³C) in plant matter. Thesame traits would also seem to be relevant when breeding forhot, irrigated environments. Where additional water is not availableto the crop, higher water-use efficiency (WUE) appears to bean alternative strategy to improve crop performance. In thiscontext ¹³C constitutes a simple but reliable measure of WUE.However, in contrast to lines performing better because of increasedaccess to water, lines producing greater biomass due to superiorWUE will have lower ¹³C values. WUE may be modified not onlythrough a decrease in stomatal conductance, but also throughan increase in photosynthetic capacity. Harvest index is stronglyreduced by terminal drought (i.e. drought during grain filling).Thus, phenological traits increasing the relative amount ofwater used during grain filling, or adjusting the crop cycleto the seasonal pattern of rainfall may be useful. Augmentingthe contribution of carbohydrate reserves accumulated duringvegetative growth to grain filling may also be worthwhile inharsh environments. Alternatively, extending the duration ofstem elongation without changing the timing of anthesis wouldincrease the number of grains per spike and the harvest indexwithout changing the amount of water utilized by the crop.     

Different acclimatization mechanisms of two grass pea cultivars to osmotic stress in in vitro culture

Grass pea (Lathyrus sativus L.) is a legume crop known from its tolerance to various abiotic stresses, especially drought. In this study, we investigated: (1) the response of grass pea seedlings to osmotic stress generated in vitro by polyethylene glycol (PEG); (2) potential drought acclimatization mechanisms of two polish grass pea cultivars. Grass pea seeds of two cultivars were sown on media containing different PEG concentrations (0, 5.5, 11.0 mM) and cultivated for 14 days in controlled conditions. Plants’ dry matter increased under osmotic stress (regardless of PEG concentration). In turn, the highest dose of PEG caused a reduction in seedling growth in both cultivars. Furthermore, PEG caused the peroxidase activity increase in whole seedlings and catalase (CAT) activity in roots. However, differences between cultivars were noted in: CAT activity in shoots; while phenols and anthocyanin content as well as electrolyte leakage in shoots and roots. In turn, in both tested genotypes, accumulation of proline increased in shoots under osmotic stress. Obtained results indicate that the examined plants, although belonging to the same species, differ in acclimatization processes leading to elevated tolerance to osmotic stress.     

Seed vigour contributes to yield improvement in dry direct‐seeded rainfed lowland rice

Dry direct‐seeded rice (DSR) cultivation is widely spreading in tropical Asia, but drought and nutrient deficiency stresses often cause crop failure in rainfed lowlands. The objective of this study was to dissect the physio‐morphological characteristics associated with crop establishment and early vigour of DSR under drought and P deficiency conditions in the Philippines. It was found that new drought‐resistant cultivars bred for DSR (Rc348 and Rc192) had faster germination and sprout growth than popular irrigated rice cultivars (Rc222 and Rc10) under soil water deficit due to rapid moisture acquisition by the germinating seeds from drying soils. There was a significant correlation between seed moisture content and the reduction in seed dry weight, and between reduction in seed dry weight and shoot elongation under both control and drought stress treatments at the germination stage. At the seedling stage, the root growth of Rc348 under drought tended to be more vigorous with its higher root‐to‐shoot ratio compared to Rc222 and Rc10. The seedling vigour of Rc348 under P deficiency was also greater than that of Rc222 due to its greater root growth and P uptake. The yields of Rc348 and Rc192 grown under rainfed condition at the target drought‐prone site where a dry spell of 13 days occurred during crop establishment were higher (4.0–4.1 t ha^?1) than the yield of Rc10 (3.0 t ha^?1). These results suggest that quick germination and seedling vigour with quick root anchorage and great nutrient uptake capacity, even with limitations of soil moisture and nutrients, would be important traits for DSR in rainfed lowlands.     

Differential accumulation of a 24-kd dehydrin protein in wheat seedlings correlates with drought stress tolerance at grain filling

The Pacific Northwest (PNW), an important region for wheat production in the USA, is often subject to water deficits during sowing and grain filling. These deficits reduce the quality and yield of the crop. As a consequence, an important objective of breeding programs in the region is improving the genetic adaptation of wheat cultivars to drought stress. One response to dehydrative stresses is the accumulation of proteins called dehydrins, which are believed to protect membranes and macromolecules against denaturation. We characterized dehydrin accumulation in seedlings during drought stress and its correlation with stress tolerance during grain filling in seven wheat cultivars, 'Connie', 'Gene', 'TAM105', 'Rod', 'Hiller', 'Rhode', and 'Stephens'. A 24-kd dehydrin accumulated in seedlings under stress, but not in irrigated control plants. Connie, TAM105, and Gene started to accumulate dehydrins at the fourth day of stress, while the other cultivars showed dehydrins after twelve days of stress. This differential accumulation in seedlings was associated with stress tolerance at grain filling, characterized by a lower reduction in yield and in the rate of decrease in leaf water potential per day of stress. Connie, TAM105, Gene and Rod where the most tolerant cultivars. The results indicate that expression of this 24-kd dehydrin might serve as a rapid and non-destructive screening technique at the seedling stage. Even though the results are promising, selection experiments using a population segregating for stress tolerance are needed to test more conclusively whether this dehydrin can serve as a genetic marker for cultivars with tolerance to drought stress.     

Crop responses to drought and the interpretation of adaptation

Drought is a multidimensional stress affecting plants at various levels of their organization. The effect of and plant response to drought at the whole plant and crop level is most complex because it reflects the integration of stress effects and responses at all underlying levels of organization over space and time. This review discusses some of the major aspects of crop response to drought stress which are relevant for plant breeding. Emphasis is given to whole plant aspects which are too often disregarded when conclusions are drawn from molecular studies towards the genetic improvement of crop drought resistance. Topics discussed are seedling emergence and establishment, plant phenology, leaf area, water deficit and assimilation, osmotic adjustment, the root and the formation of yield. The discussion is concluded with the interpretation of crop adaptation to drought conditions in its agronomic sense. Conclusions are drawn regarding plant breeding for drought-prone conditions.     

东乡普通野生稻与栽培稻苗期抗旱性的比较

干旱影响水稻生长发育,不论什么时期发生最终都导致产量损失。研究水稻资源抗旱性有助于水稻抗旱改良和稳定干旱胁迫下水稻的产量。东乡普通野生稻被公认为是栽培稻的祖先,对增强水稻抗旱性可能十分重要。对4份来自3个仅存的居群的东乡野生稻与15份栽培稻进行苗期抗旱性比较,考察了3次重复的盆栽土培试验中8个抗旱指标。表明东乡普通野生稻比栽培稻更为抗旱,表现在最大根长、茎长、根干重、根鲜重、根干鲜重比及抗旱指数等6个性状,而不表现在根数及根茎长比;其中茎长、最长根长、根干重、根鲜重及根系相对含水量对水稻苗期抗旱性影响更大。采用抗旱指数和抗旱总级别值法对水稻抗旱性进行评定,结果表明4份东乡野生稻材料间的抗旱性存在很大差异,且来水桃树下居群的抗性最高,东乡野生稻抗旱性可能与其原生境状况有关。结果认为东乡普通野生稻可作为栽培稻抗旱改良的遗传资源。     

Analysis of early responses to drought associated with field drought adaptation in four Sahelian groundnut (Arachis hypogaea L.) cultivars

Groundnut (Arachis hypogaea L.) is the most important oil and cash crop in the sub-Sahelian tropics. Plant adaptation to drought, i.e. cultivars (cvs) that can maintain yield when water is limited, is a complex phenomenon which is not yet fully understood. This study aimed to identify traits expressed at the early stages of the cycle that could reveal cv differences in drought adaptation in the field. The field productivity of four Sahelian groundnut cvs was assessed during three crop seasons in Bambey (Senegal). The same cvs grown in rhizotrons were subjected to early drought stress and to a desiccation test to assess cell membrane tolerance. Between-cv differences were found with respect to pod yield, biomass production, water use efficiency (WUE), stomatal regulation and cell membrane tolerance. Two strategies to cope with water deficit were identified. The first behaviour was characterised by high rapid water loss, late stomatal closure and low cell membrane damage during drought. These traits are all found in the semi-late Virginia cv 57-422 and, into a lesser extent, in the early Spanish cv Fleur 11. For both cvs, biomass production was boosted under favourable conditions in rhizotrons but the semi-late cv had poor pod yield under end-of-season water deficit conditions. The second strategy involved opposite characters, leading to the maintenance of a higher water status, resulting in lower photosynthesis and yield. This characterised the early Spanish cv 73-30, and also, to some extent, the early Spanish cv 55-437. Earliness associated with high WUE, stomatal conductance and cell membrane tolerance, were the main traits of Fleur 11, a cv derived from a Virginia × Spanish cross, which was able to maintain acceptable yield under varying drought patterns in the field. These traits, as they were detectable at an early stage, could therefore be efficiently integrated in groundnut breeding programmes for drought adaptation.