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.     

Pigeonpea breeding in eastern and southern Africa: challenges and opportunities

Pigeonpea (Cajanus cajan [L.] Millspaugh) is an important multipurpose grain legume crop primarily grown in tropical and subtropical areas of Asia, Africa and Latin America. In Africa, the crop is grown for several purposes including food security, income generation, livestock feed and in agroforestry. Production in Eastern and Southern Africa (ESA) is however faced with many challenges including limited use of high‐yielding cultivars, diseases and pests, drought, under‐investment in research and lack of scientific expertise. The aim of this review is to highlight the challenges facing pigeonpea breeding research in ESA and the existing opportunities for improving the overall pigeonpea subsector in the region. We discuss the potential of the recently available pigeonpea genomic resources for accelerated molecular breeding, the prospects for conventional breeding and commercial hybrid pigeonpea, and the relevant seed policies, among others, which are viewed as opportunities to enhance pigeonpea productivity.     

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.     

Is Discrimination of 13C in Potato Leaflets and Tubers an Appropriate Trait to Describe Genotype Responses to Restrictive and Well‐Watered Conditions?

Selection for drought tolerance entails prioritizing plant traits that integrate critical physiological processes occurring during crop growth. Discrimination against ¹³C (?) in leaflets (?_leaflet) and tubers (?_tuber) was compared under two water regimes in two potato‐improved varieties selected to maintain yield under drought conditions (Unica and Sarnav) and one drought susceptible European cultivar (Désirée). In the control treatment, soil water content was kept at field capacity over the whole growth cycle, while in the drought treatment water supply was restricted after tuber initiation (50 % of field capacity). Gas exchange and N content per unit leaf area (N_area) as well as ? were assessed at different stages. Sarnav showed the highest tuber yield in both water conditions, suggesting that yield in the water restriction treatment was largely driven by yield potential in this genotype. Higher stomatal conductance (g_s) and N_area and lower ?_leaflet in well‐watered Sarnav suggested higher photosynthetic capacity. Under water restriction, Sarnav maintained higher g_s indicating that carbon diffusion was a key factor for biomass accumulation under water restriction. Our results suggest the use of ? determined after tuber initiation as an indirect selection indicator for tuber yield under both well‐watered and restricted soil water availability conditions.     

Fulvic Acid Application Improves the Maize Performance under Well‐watered and Drought Conditions

Water deficit is perhaps the most severe threat to sustainable crop production in the conditions of changing climate. Researchers are striving hard to develop resistance against water deficit in crop plants to ensure food security for the coming generations. This study was conducted to establish the role of fulvic acid (FA) application in improving the performance of hybrid maize (Zea mays L.) under drought. Maize plants were grown under normal conditions till tasselling and were then subjected to drought by cessation of water followed by foliar application of FA (1.5 mg l^?1). Drought stress disrupted the photosynthetic pigments and reduced the gas exchange leading to reduction in plant growth and productivity. Nonetheless, exogenous FA application substantially ameliorated the adversities of drought by sustaining the chlorophyll contents and gas exchange possibly by enhanced levels of antioxidant enzyme (superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT)) activities and proline. These beneficial effects yielded in terms of plant growth and allometry, and grain yield. It is interesting to note that FA application also improved the crop performance under well‐watered conditions. Hence, FA may be applied to improve the crop performance under drought and well‐watered conditions.     

Forage legumes for future dry climates: Lower relative biomass losses of minor forage legumes compared to Trifolium repens under conditions of periodic drought stress

Dairy livestock production systems rely on high‐quality forage legumes, which are widely present in grassland swards all over Europe. A future climatic scenario with higher average annual temperatures and lower precipitation is expected to affect grassland productivity in general and the productivity of the most important forage legume species Trifolium repens in particular. One way to cope with such constraints is the adoption of currently underutilized minor legume species with a higher tolerance towards drought stress. Therefore, the present study investigated legume species with lower moisture requirements than T. repens, these are Lotus corniculatus, Medicago falcata, Medicago lupulina and Onobrychis viciifolia in comparison with T. repens. Legumes were grown in containers as monocultures in a roofed open‐sided greenhouse under conditions of optimal water supply or periodic drought stress. Generally, drought stress decreased the biomass production, but species differed in their reaction. Particularly, M. lupulina and L. corniculatus had lower relative biomass losses (?26%) compared to T. repens (?43%, as averaged over drought stress periods). However, in overall biomass production T. repens still was at one level with M. lupulina and L. corniculatus under drought stress. This was related to high stomata control as indicated by the intrinsic water‐use efficiency. We conclude that there are promising future options of forage legumes as alternatives to T. repens.     

Effect of Simulating Drought in Various Phenophases on the Water Use Efficiency of Winter Wheat

In Central Europe, drought is the most important limiting factor for autumn‐sown cereals. Due to the decline in groundwater, it is a priority to use less water‐demanding forms of crop production. Water use efficiency (WUE) can only be increased if cultivars with satisfactory water management traits are grown, so that they can exploit the water reserves of the soil even if drought occurs during the vegetation. Water consumption and water use efficiency of winter wheat genotypes were investigated in a model experiment carried out in a climate‐controlled glasshouse. The plants were grown either with optimum water supplies or with simulated drought in three phenophases, and measurements were made on the yield parameters, phenological traits and water use parameters of the plants. Substantial differences were observed between the water demands of the cultivars, and it was found that the later the phenophase in which drought was simulated, the greater the decline in water uptake. The analysis of WUE led to the conclusion that the WUE values of cultivars with short vegetation periods dropped to the greatest extent when water deficit was suffered at first node appearance, while cultivars with longer vegetation periods were more sensitive to drought during the heading and grain‐filling stages.     

Wheat Rht‐B1 Dwarfs Exhibit Better Photosynthetic Response to Water Deficit at Seedling Stage Compared to the Wild Type

Wheat reduced height (Rht) genes encode modified DELLA proteins, which are gibberellin insensitive, accumulate under stress, restrain growth and affect plant stress response. The seedling reaction to soil water deficit regarding leaf gas exchange and chlorophyll fluorescence was compared in near‐isogenic lines carrying the alleles Rht‐B1a (tall), Rht‐B1b (semi‐dwarfing) and Rht‐B1c (dwarfing) and was related to leaf water content and anatomy. Under drought, Rht‐B1c line was characterized by less decreased CO₂ assimilation, delayed non‐stomatal limitation of photosynthesis and higher instantaneous water use efficiency. The functional state of its photosynthetic apparatus was better preserved as evidenced by the less decreased actual quantum yield (Φ_PSII) and potential maximum quantum yield (F_v/F_m) of PSII, and the less increased quantum yield of non‐regulated energy dissipation (Φ_NO). Rht‐B1b line also tended to perform better than Rht‐B1a, but differences were less pronounced. Although the leaves of both dwarf lines were smaller, thicker and more pubescent, their water content was not higher in comparison with the tall line. Nevertheless, in Rht‐B1c, leaf thickness was less decreased and mesophyll cells were less shrunk under drought. The more effective performance of the photosynthetic machinery of dwarf lines under water deficit could be explained by a combination of morpho‐anatomical and metabolic characteristics.     

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.     

Phosphorus Fertilization and Fungal Inoculations Affected the Physiology,Phosphorus Uptake and Growth of Spring Wheat Under Rainfed Conditions on the Canadian Prairies

Some fungal species have been shown to improve plant growth under drought conditions and to increase plant phosphorus (P) uptake from the soil. How moisture limitation, P availability and fungal inoculation interact to affect plant physiology and growth is, however, poorly understood. Here, we studied the combined effects of fungal (arbuscular mycorrhizal fungi (AMF) or Penicillium spp.) inoculations and phosphorus (P) fertilization (0, 45 and 90 kg ha^?1) on the net rate of photosynthesis, water‐use efficiency, P uptake and growth of spring wheat (Triticum aestivum var. Superb) under field conditions at two locations (Castor and Vegreville) in Alberta, Canada. Both fungal inoculation and P application increased the rate of photosynthesis. Under the same P level, AMF inoculation had a greater positive effect on the rate of photosynthesis than Penicillium inoculation. The AMF inoculation increased the instantaneous water‐use efficiency (WUEi) of plants at Castor, but not at Vegreville. Leaf carbon isotope discrimination (CID, Δ¹³C) increased with the rate of P application but was not affected by fungal inoculations. Phosphorus concentrations of stem and seed increased with both fungal inoculation and P application irrespective of location, with AMF inoculation showing the largest effects. The interaction between P addition and fungal inoculation was significant for stem P concentration in Vegreville. Both fungal inoculation and P application increased the leaf area index (LAI), biomass production and grain yield at both locations. Under the same P level, AMF inoculation had a greater positive effect on LAI, biomass production and grain yields than Penicillium inoculation. Morphological characters such as spike length and kernels/spike were also improved by fungal inoculation and P application at both locations. We conclude that the studied sites were deficient in P availability, and both fungal inoculation and P application improved P uptake and crop productivity, while the effect of fungal inoculation on water‐use efficiency was site specific.     

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.     

Genetic diversity of tropical maize inbred lines combining resistance to Striga hermonthica with drought tolerance using SNP markers

Striga hermonthica and drought are the major stresses limiting maize yields in sub‐Saharan Africa. The search for diverse maize lines’ tolerance to drought and resistance to S. hermonthica (DTSTHR) is very crucial for yield improvement in areas affected by the two stresses. Understanding the genetic diversity among the lines is important to develop cultivars resistant to S. hermonthica and tolerant to drought. The lines were developed from biparental crosses of drought‐tolerant and Striga‐resistant lines. A total of 128 DTSTHR maize lines were characterized using single‐nucleotide polymorphism (SNP) markers. Results of the cluster analysis based on 3297 SNP markers showed four distinct groups consistent with the pedigrees of the lines. Furthermore, model‐based analysis also formed the same groups of the DTSTHR lines. Integrating the pedigree information with combining ability and the SNP analyses may provide defined heterotic groups for maize improvement work in West and Central Africa. These results also help breeders to utilize DTSTHR lines present at IITA for developing biparental crosses without disrupting the heterotic groups they have established in their breeding programmes.     

Nitrogen Sustains Seed Yield of Quinoa Under Intermediate Drought

Quinoa (Chenopodium quinoa Willd.) is a promising crop for food security in dry areas. Studies have been conducted to define nitrogen (N) fertilization levels and to understand the responses of quinoa to drought, but little is known about the response of this crop to N fertilization under drought stress. The aim of this study was to investigate whether N fertilization could improve quinoa yield and physiology under limited water. A greenhouse experiment was carried out with quinoa grown at four N fertilization levels (0, 0.2, 0.4 and 0.6 g N pot^?1) and two watering treatments (progressive drought and full irrigation; 10 and 98 % of pot water holding capacity, respectively). Results of this experiment showed that N may confer a certain degree of drought tolerance to quinoa as seed quality and yield of N‐fertilized plants were not affected by drought stress. Responses such as faster stomatal closure, reduced leaf water potential, higher leaf abscisic acid (ABA) concentration and particularly an improved N remobilization in N‐fertilized plants may have played a role in sustaining seed yield in the drought‐stressed treatment. These results under controlled conditions serve as a basis to elucidate drought tolerance mechanisms activated with N fertilization and to define the use of N in management practices under semi‐arid environments.     

Sugarcane Breeding Institute,Coimbatore, India Effect of Different Levels of Drought during the Formative Phase on Growth Parameters and its Relationship with Dry Matter Accumulation in Sugarcane

A field experiment was conducted at Coimbatore (11° N, 77° E), India during the 1996 and 1997 crop seasons, using four commercial sugarcane varieties (Co 8021, Co 419, Co 8208 and Co 6304), to study the effect of three levels of drought (severe, moderate and control) during the formative phase (60–150 days after planting) on growth determinants and their relationship with dry matter accumulation. The reduction in dry matter content was 60.8, 52.4 and 25.9 % in severe drought and 46.3, 36.3 and 15.1 % in moderate drought at the ends of the formative, grand growth and maturity phases, respectively. High net assimilation rate, optimum leaf area index, high crop growth rate and an early shift in dry matter allocation to the stem were found to be desirable for higher biomass production, especially under water‐limited drought conditions. Measurement of growth parameters such as net assimilation rate, relative growth rate, leaf area index and leaf area duration under drought and crop growth rate and stalk elongation rate under normal irrigated conditions, particularly during the formative phase, might help to predict total dry matter at harvest. Leaf area ratio was not found to correlate with total dry matter at harvest in either drought or normal irrigated conditions.     

Drought Tolerance and Water Use of Cereal Crops: A Focus on Sorghum as a Food Security Crop in Sub‐Saharan Africa

Sub‐Saharan Africa (SSA) faces twin challenges of water stress and food insecurity – challenges that are already pressing and are projected to grow. Sub‐Saharan Africa comprises 43 % arid and semi‐arid area, which is projected to increase due to climate change. Small‐scale, rainfed agriculture is the main livelihood source in arid and semi‐arid areas of SSA. Because rainfed agriculture constitutes more than 95 % of agricultural land use, water scarcity is a major limitation to production. Crop production, specifically staple cereal crop production, will have to adapt to water scarcity and improved water productivity (output per water input) to meet food requirements. We propose inclusion and promotion of drought‐tolerant cereal crops in arid and semi‐arid agro‐ecological zones of SSA where water scarcity is a major limitation to cereal production. Sorghum uniquely fits production in such regions, due to high and stable water‐use efficiency, drought and heat tolerance, high germplasm variability, comparative nutritional value and existing food value chain in SSA. However, sorghum is socio‐economically and geographically underutilized in parts of SSA. Sorghum inclusion and/or promotion in arid and semi‐arid areas of SSA, especially among subsistence farmers, will improve water productivity and food security.     

转ppc基因水稻苗期抗旱特性研究

通过C₄转基因技术改善C₃作物光合作用, 以期提高作物产量是国内外研究的热点之一。然而, 目前关于转磷酸烯醇式丙酮酸羧化酶(PEPC)基因对水稻的光合作用、产量和抗旱性的影响及其调节机理仍不很清楚。本研究以T₄代转ppc基因水稻为材料, 进行产量和苗期抗旱性研究。结果表明, 在旱作栽培条件下, 两个转ppc基因株系(T1, T2)单株产量分别比未转基因的对照(WT)增产28%和42%, 分蘖增加27%和40%; T1和T2可维持较高的光合速率、气孔导度和水分利用效率; 干旱胁迫使超氧化物歧化酶(SOD)活性和丙二醛(MDA)含量增加, T1和T2 SOD活性增幅(+25%)都显著高于WT(+9%), 而MDA含量增幅显著低于WT, 表明转ppc基因水稻具有较强的抗氧化能力; T1和T2比WT具有较高的脯氨酸含量和渗透势下降幅度, 说明转PEPC基因水稻的渗透调节能力高于对照。PEG-6000处理下, 得到的结果相似。     

Phenotypic Changes in Grain Sorghum Over the Last Five Decades

Sorghum [Sorghum bicolor (L.) Moench] is a drought‐tolerant crop, and its productivity in rain fed environments has increased since the 1950s. This increase is due to changes in agronomic practices and hybrid improvement. The objective of this study was to determine what aspects of grain sorghum morphology, physiology and water use have changed with hybrid improvement and might have contributed to this yield increase. A 2‐year greenhouse experiment was conducted with one hybrid from each of the past five decades. The hybrids were studied in well‐watered and pre‐ and post‐flowering water deficit conditions. Total water use, transpiration, stomatal conductance and photosynthesis were measured during the growing period. Biomass and biomass components were measured at harvest. There was no consistent change in the leaf physiological parameters resulting from hybrid advancement. In contrast, total water use increased in rate of 8.5 cm³ kg soil^?1 year^?1 from old to new hybrids in the well‐watered treatments. Root biomass also increased in rate of 0.2 g plant^?1 year^?1. Leaf biomass and panicle length also was greater for the newest compared with the older hybrids. Hybrid advancement was related to increase in panicle length but decrease in peduncle length. Results indicated that hybrid development programmes created hybrids with improved drought avoidance, due to better root density of newly released hybrids, or hybrids with better resource use which ultimately increased yield under rain fed conditions.     

Mild Drought Stress‐Induced Changes in Yield,Physiological Processes and Chemical Composition in Festuca,Lolium and Festulolium

The impact of mild drought stress (3 weeks at 40 % field water capacity) on yield, physiological processes, accumulation of proline and phenolic compounds and forage quality parameters in forage grasses was evaluated in pot experiments. During four different growing periods, the effects of water deficit were assessed in nine varieties from five species (Lolium perenne, Lolium multiflorum, Festuca pratensis, Festuca arundinacea and Festulolium braunii). All measured parameters were affected by drought stress in the different cuts. Photosynthesis, transpiration rate, stomatal conductance and dry matter yield were significantly lower in drought stress than under well‐watered conditions in all varieties. Higher water‐use efficiency was only observed during the first and fourth drought period, while this was not the case in the second and third. Mild drought stress significantly increased the content of proline, phenolic acid, flavonoids, water‐soluble carbohydrates and protein. All tested grasses showed also an increase of organic matter digestibility and cell wall digestibility under drought stress conditions.     

Effects of Salinity and Soil–Drying on Radiation Use Efficiency,Water Productivity and Yield of Quinoa (Chenopodium quinoa Willd.)

Drought and salinity reduce crop productivity especially in arid and semi‐arid regions, and finding a crop which produces yield under these adverse conditions is therefore very important. Quinoa (Chenopodium quinoa Willd.) is such a crop. Hence, a study was conducted in field lysimeters to investigate the effect of salinity and soil–drying on radiation use efficiency, yield and water productivity of quinoa. Quinoa was exposed to five salinity levels (0, 10, 20, 30 and 40 dS m^?1) of irrigation water from flower initiation onwards. During the seed‐filling phase the five salinity levels were divided between two levels of irrigation, either full irrigation (FI; 95 % of field capacity) or non‐irrigated progressive drought (PD). The intercepted photosynthetically active radiation was hardly affected by salinity (8 % decrease at 40 dS m^?1) and did not differ significantly between FI and PD. Radiation use efficiency of dry matter was similar between salinity levels and between FI and PD. In line with this, no negative effect of severe salinity and soil–drying on total dry matter could be detected. Salinity levels between 20 and 40 dS m^?1 significantly reduced the seed yield by ca. 33 % compared with 0 dS m^?1 treatment owing to a 15–30 % reduction in seed number per m², whereas the seed yield of PD was 8 % less than FI. Consequently, nitrogen harvested in seed was decreased by salinity although the total N‐uptake was increased. Both salinity and drought increased the water productivity of dry matter. Increasing salinity from 20 to 40 dS m^?1 did not further decrease the seed number per m² and seed yield, which shows that quinoa (cv. Titicaca) acclimated to saline conditions when exposed to salinity levels between 20 and 40 dS m^?1.     

Improving adaptation to drought stress in white pea bean (Phaseolus vulgaris L.): Genotypic effects on grain yield,yield components and pod harvest index

Common bean (Phaseolus vulgaris L.) is the most important food legume crop in Africa and Latin America where rainfall pattern is unpredictable. The objectives were to identify better yielding common bean lines with good canning quality under drought, and to identify traits that could be used as selection criteria for evaluating drought‐tolerant genotypes. In all, 35 advanced lines were developed through single seed descent and evaluated with a standard check under drought and irrigated conditions at two locations over 2 years in Ethiopia. Grain yield (GY), pod number per m², seed number per m² and seed weight decreased by 56%, 47%, 49% and 14%, respectively, under drought stress. Eight genotypes had better yield with good canning quality under drought compared to the check. Moderate to high proportion of genetic effects were observed under drought conditions for GY and yield components compared to genotype × environment effects. Significant positive correlations between GY and pod harvest index (PHI) in drought suggest that PHI could be used as an indirect selection criterion for common bean improvement.