Plant breeding and drought in C3 cereals: what should we breed for?
ABSTRACT: Drought is the main abiotic constraint on cereal yield. Analysing physiological determinants of yield responses to water may help in breeding for higher yield and stability under drought conditions. The traits to select (either for stress escape, avoidance or tolerance) and the framework where breeding for drought stress is addressed will depend on the level and timing of stress in the targeted area. If the stress is severe, breeding under stress-free conditions may be unsuccessful and traits that confer survival may become a priority. However, selecting for yield itself under stress-alleviated conditions appears to produce superior cultivars, not only for optimum environments, but also for those characterized by frequent mild and moderate stress conditions. This implies that broad avoidance/tolerance to mild-moderate stresses is given by constitutive traits also expressed under stress-free conditions. In this paper, we focus on physiological traits that contribute to improved productivity under mild-moderate drought. Increased crop performance may be achieved through improvements in water use, water-use efficiency and harvest index. The first factor is relevant when soil water remains available at maturity or when deep-rooted genotypes access water in the soil profile that is not normally available; the two latter conditions become more important when all available water is exhausted by the end of the crop cycle. Independent of the mechanism operating, a canopy able to use more water than another would have more open stomata and therefore higher canopy temperature depression, and 13C discrimination (delta13C) in plant matter. The same traits would also seem to be relevant when breeding for hot, irrigated environments. Where additional water is not available to the crop, higher water-use efficiency (WUE) appears to be an alternative strategy to improve crop performance. In this context delta13C constitutes a simple but reliable measure of WUE. However, in contrast to lines performing better because of increased access to water, lines producing greater biomass due to superior WUE will have lower delta13C values. WUE may be modified not only through a decrease in stomatal conductance, but also through an increase in photosynthetic capacity. Harvest index is strongly reduced by terminal drought (i.e. drought during grain filling). Thus, phenological traits increasing the relative amount of water used during grain filling, or adjusting the crop cycle to the seasonal pattern of rainfall may be useful. Augmenting the contribution of carbohydrate reserves accumulated during vegetative growth to grain filling may also be worthwhile in harsh environmcnts. Alternatively, extending the duration of stem elongation without changing the timing of anthesis would increase the number of grains per spike and the harvest index without changing the amount of water utilized by the crop.
Project description:High water use efficiency (WUE) can be achieved by coordination of biomass accumulation and water consumption. WUE is physiologically and genetically linked to carbon isotope discrimination (CID) in leaves of plants. A population of 148 recombinant inbred lines (RILs) of sunflower derived from a cross between XRQ and PSC8 lines was studied to identify quantitative trait loci (QTL) controlling WUE and CID, and to compare QTL associated with these traits in different drought scenarios. We conducted greenhouse experiments in 2011 and 2012 by using 100 balances which provided a daily measurement of water transpired, and we determined WUE, CID, biomass and cumulative water transpired by plants. Wide phenotypic variability, significant genotypic effects, and significant negative correlations between WUE and CID were observed in both experiments. A total of nine QTL controlling WUE and eight controlling CID were identified across the two experiments. A QTL for phenotypic response controlling WUE and CID was also significantly identified. The QTL for WUE were specific to the drought scenarios, whereas the QTL for CID were independent of the drought scenarios and could be found in all the experiments. Our results showed that the stable genomic regions controlling CID were located on the linkage groups 06 and 13 (LG06 and LG13). Three QTL for CID were co-localized with the QTL for WUE, biomass and cumulative water transpired. We found that CID and WUE are highly correlated and have common genetic control. Interestingly, the genetic control of these traits showed an interaction with the environment (between the two drought scenarios and control conditions). Our results open a way for breeding higher WUE by using CID and marker-assisted approaches and therefore help to maintain the stability of sunflower crop production.
Project description:Drought stress has been identified as the major environmental factor limiting soybean [Glycine max (L.) Merr.] yield worldwide. Current breeding efforts in soybean largely focus on identifying genotypes with high seed yield and drought tolerance. Water use efficiency (WUE) that results in greater yield per unit rainfall is an important parameter in determining crop yields in many production systems, and is often related with crop drought tolerance. Even though roots are major plant organs that perceive and respond to drought stress, their utility in improving soybean yield and WUE under different environmental and management conditions are largely unclear. The objectives of this research was to evaluate soybean cultivars and breeding and germplasm lines for yield, WUE, root penetrability of hardpan, and root morphology. Field experiments were conducted at two locations in South Carolina (southeastern United States) during the 2017 cropping season to test the genotypes for yield and root morphology under irrigated and non-irrigated conditions. Two independent controlled-environmental experiments were conducted to test the genotypes for WUE and root penetrability of synthetic hardpans. The slow wilting lines NTCPR94-5157 and N09-13890 had equal or greater yield than the checks- cultivar NC-Raleigh and the elite South Carolina breeding line SC07-1518RR, under irrigated and non-irrigated conditions. The high yielding genotypes NTCPR94-5157, N09-13890, and SC07-1518RR exhibited root parsimony (reduced root development). This supported the recent hypothesis in literature that root parsimony would have adaptational advantage to improve yield under high input field conditions. The high yielding genotypes NTCPR94-5157, N09-13890, NC-Raleigh, and SC07-1518RR and a cultivar Boggs (intermediate in yield) possessed high WUE and had increased root penetrability of hardpans. These genotypes offer useful genetic materials for soybean breeding programs for improving yield, drought tolerance, and/or hardpan penetrability.
Project description:Trait-based breeding is essential to improve wheat yield, particularly when stress adaptation is targeted. A set of modern and underutilized wheat genotypes was examined in a 2-year field experiment with distinct seasonal water supply. Yield formation and drought response strategies were analyzed in relation to components of Passioura's yield-water framework based on phenological, morphological, physiological, and root characteristics. Limited water supply resulted in 60% yield loss and substantially lower water use (37%), water use efficiency (32.6%), and harvest index (14%). Phenology and root length density were key determinants of water use. Late flowering underutilized wheat species with large root system and swift ground coverage showed greatest water use. Leaf chlorophyll concentration and stomata conductance were higher in modern cultivars, supporting their high biomass growth and superior water use efficiency. While, lower chlorophyll concentration and stomata conductance of underutilized wheats indicated a water saving strategy with an intrinsic limitation of potential growth. Harvest index was strongly dependent on phenology and yield components. Optimized flowering time, reduced tillering, and strong grain sink of modern cultivars explained higher harvest index compared to underutilized wheats. Cluster analysis revealed the consistent differentiation of underutilized and modern wheats based on traits underlying Passioura's yield-water framework. We identified physiological and root traits within modern cultivars to be targeted for trait-based crop improvement under water-limited conditions. High capacity of water use in underutilized genetic resources is related to yield-limiting phenological and morphological traits, constraining their potential role for better drought resistance. Still some genetic resources provide adaptive features for stress resistance compatible with high yield as revealed by high harvest index under drought of Khorasan wheat.
Project description:Increasing crop productivity under conditions of climate change requires the identification, selection, and utilization of novel alleles for breeding. In this study, we analysed the genotype and field phenotype of the barley HEB-25 multi-parent mapping population under well-watered and water-limited environments for two years. A genome-wide association study (GWAS) for genotype × environment interactions was performed for 10 traits including flowering time (heading time, HEA) and plant grain yield (PGY). Comparison of the GWAS for traits per se (i.e. regardless of the environment) with a study for quantitative trait loci (QTLs) × environment interactions (Q×E), indicates the prevalence of Q×E mostly for reproductive traits. One Q×E locus on chromosome 2, Hordeum spontaneum Dry2.2 (HsDry2.2), showed a positive and conditional effect on PGY and grain number (GN). The wild allele significantly reduced HEA; however, this earliness was not conditioned by water deficit. Furthermore, BC2F1 lines segregating for the HsDry2.2 locus showed that the wild allele conferred an advantage over the cultivated allele in PGY, GN, and harvest index, as well as modified shoot morphology, a longer grain-filling period, and reduced senescence (only under drought). This suggests the presence of an adaptation mechanism against water deficit rather than an escape mechanism. The study highlights the value of evaluating wild relatives in search of novel alleles and provides clues to resilience mechanisms underlying crop adaptations to abiotic stress.
Project description:End-of-season drought or "terminal drought," which occurs after flowering, is considered the most significant abiotic stress affecting crop yields. Wheat crop production in Mediterranean-type environments is often exposed to terminal drought due to decreasing rainfall and rapid increases in temperature and evapotranspiration during spring when wheat crops enter the reproductive stage. Under such conditions, every millimeter of extra soil water extracted by the roots benefits grain filling and yield and improves water use efficiency (WUE). When terminal drought develops, soil dries from the top, exposing the top part of the root system to dry soil while the bottom part is in contact with available soil water. Plant roots sense the drying soil and produce signals, which on transmission to shoots trigger stomatal closure to regulate crop water use through transpiration. However, transpiration is linked to crop growth and productivity and limiting transpiration may reduce potential yield. While an early and high degree of stomatal closure affects photosynthesis and hence biomass production, a late and low degree of stomatal closure exhausts available soil water rapidly which results in yield losses through a reduction in post-anthesis water use. The plant hormone abscisic acid (ABA) is considered the major chemical signal involved in stomatal regulation. Wheat genotypes differ in their ability to produce ABA under drought and also in their stomatal sensitivity to ABA. In this viewpoint article we discuss the possibilities of exploiting genotypic differences in ABA response to soil drying in regulating the use of water under terminal drought. Root density distribution in the upper drying layers of the soil profile is identified as a candidate trait that can affect ABA accumulation and subsequent stomatal closure. We also examine whether leaf ABA can be designated as a surrogate characteristic for improved WUE in wheat to sustain grain yield under terminal drought. Ease of collecting leaf samples to quantify ABA compared to extracting xylem sap will facilitate rapid screening of a large number of germplasm for drought tolerance.
Project description:Drought is one of the major abiotic stresses affecting wheat yield. A recurrent selection program was conducted to improve the drought tolerance and yield of bread wheat using drought tolerant advanced breeding lines from a drought tolerant x susceptible cross (HI 1500 x HUW 510). The parental lines were evaluated for yield, biomass and harvest index (HI) in addition to the drought adaptive traits like Canopy Temperature (CT), chlorophyll content and Normalized Difference Vegetative Index (NDVI). After three rounds of recurrent selection, the half sib progenies exhibited a marked reduction in CT, chlorophyll content and biomass; whereas improvement was noted for yield, HI and NDVI. Drought tolerance of the half-sib population appeared enhanced as indicated by drought indices and grain yield. Compared to base population, half-sibs showed better HI, grain filling and a significant (17.1%) increase in grain yield under water stress conditions. Cooler canopies and increased early vigour might have contributed to drought tolerance. A favourable combination of gibberellin sensitive and insensitive Rhtalleles was observed in the recombinant progenies. However, increased yield under water stress had a negative trade off in reduction of biomass. The study also identified potential lines with high yield and drought tolerance for subsequent varietal development for water limited areas.
Project description:Water deficit is a serious environmental factor limiting the growth and productivity of plants worldwide. Improvement of drought tolerance and efficient water use are significant strategies to overcome this dilemma. In this study, a drought-responsive transcription factor, nuclear factor Y subunit B 7 (PdNF-YB7), induced by osmotic stress (PEG6000) and abscisic acid, was isolated from fast-growing poplar clone NE-19 [Populus nigra × (Populus deltoides × Populus nigra)]. Ectopic overexpression of PdNF-YB7 (oxPdB7) in Arabidopsis enhanced drought tolerance and whole-plant and instantaneous leaf water-use efficiency (WUE, the ratio of biomass produced to water consumed). Overexpressing lines had an increase in germination rate and root length and decrease in water loss and displayed higher photosynthetic rate, instantaneous leaf WUE, and leaf water potential to exhibit enhanced drought tolerance under water scarcity. Additionally, overexpression of PdNF-YB7 in Arabidopsis improved whole-plant WUE by increasing carbon assimilation and reducing transpiration with water abundance. These drought-tolerant, higher WUE transgenic Arabidopsis had earlier seedling establishment and higher biomass than controls under normal and drought conditions. In contrast, Arabidopsis mutant nf-yb3 was more sensitive to drought stress with lower WUE. However, complementation analysis indicated that complementary lines (nf-yb3/PdB7) had almost the same drought response and WUE as wild-type Col-0. Taken together, these results suggest that PdNF-YB7 positively confers drought tolerance and improves WUE in Arabidopsis; thus it could potentially be used in breeding drought-tolerant plants with increased production even under water deficiency.
Project description:Stay-green trait enhances sorghum adaptation to post-flowering drought. Six stay-green backcross introgression lines (BILs) carrying one or more stay-green QTLs (Stg1-4) and their parents were characterized under non-stress (W100: 100% of soil field capacity (FC)) and two levels of post-flowering drought (W75: 75% FC; W50: 50% FC) in a controlled condition. We aimed to study the response and identify the drought threshold of these QTLs under different levels of post-flowering drought and find traits closely contributing to grain yield (GY) under different drought severity. W50 caused the highest reduction in BILs performance. From W100 to W50, the GY of the recurrent parent reduced by 70%, whereas that of the BILs reduced by only 36%. W75 and W50 induce different behavior/response compared to W100. Harvest index contributed to the GY under the three water regimes. For high GY under drought transpiration rate at the beginning of drought and mid-grain filling was important at W75, whereas it was important at mid-grain filling and late-grain filling at W50. Stay-green trait can be scored simply with the relative number of green leaves/plants under both irrigated and stress environments. QTL pyramiding might not always be necessary to stabilize or increase the GY under post-flowering drought. The stay-green QTLs increase GY under drought by manipulating water utilization depending on drought severity.
Project description:The ability of plants to acquire and use water is critical in determining life-history traits such as growth, flowering, and allocation of biomass into reproduction. In this context, a combination of functionally linked traits is essential for plants to respond to environmental changes in a coordinated fashion to maximize resource use efficiency. We analyzed different water-use traits in Arabidopsis ecotypes to identify functionally linked traits that determine water use and plant growth performance. Water-use traits measured were (i) leaf-level water-use efficiency (WUE i ) to evaluate the amount of CO 2 fixed relative to water loss per leaf area and (ii) short-term plant water use at the vegetative stage (VWU) as a measure of whole-plant transpiration. Previously observed phenotypic variance in VWU, WUE i and life-history parameters, highlighted C24 as a valuable ecotype that combined drought tolerance, preferential reproductive biomass allocation, high WUE i , and reduced water use. We therefore screened 35 Arabidopsis ecotypes for these parameters, in order to assess whether the phenotypic combinations observed in C24 existed more widely within Arabidopsis ecotypes. All parameters were measured on a short dehydration cycle. A segmented regression analysis was carried out to evaluate the plasticity of the drought response and identified the breakpoint as a reliable measure of drought sensitivity. VWU was largely dependent on rosette area, but importantly the drought sensitivity and plasticity measures were independent of the transpiring leaf surface. A breakpoint at high rSWC indicated a more drought-sensitive plant that closed stomata early during the dehydration cycle and consequently showed stronger plasticity in leaf-level WUE i parameters. None of the sensitivity, plasticity, or water-use measurements were able to predict the overall growth performance; however, there was a general trade-off between vegetative and reproductive biomass. PCA and hierarchical clustering revealed that C24 was unique among the 35 ecotypes in uniting all the beneficial water use and stress tolerance traits, while also maintaining above average plant growth. We propose that a short dehydration cycle, measuring drought sensitivity and VWU is a fast and reliable screen for plant water use and drought response strategies.
Project description:Flowering time and water-use efficiency (WUE) are two ecological traits that are important for plant drought response. To understand the evolutionary significance of natural genetic variation in flowering time, WUE, and WUE plasticity to drought in Arabidopsis thaliana, we addressed the following questions: (1) How are ecophysiological traits genetically correlated within and between different soil moisture environments? (2) Does terminal drought select for early flowering and drought escape? (3) Is WUE plasticity to drought adaptive and/or costly? We measured a suite of ecophysiological and reproductive traits on 234 spring flowering accessions of A. thaliana grown in well-watered and season-ending soil drying treatments, and quantified patterns of genetic variation, correlation, and selection within each treatment. WUE and flowering time were consistently positively genetically correlated. WUE was correlated with WUE plasticity, but the direction changed between treatments. Selection generally favored early flowering and low WUE, with drought favoring earlier flowering significantly more than well-watered conditions. Selection for lower WUE was marginally stronger under drought. There were no net fitness costs of WUE plasticity. WUE plasticity (per se) was globally neutral, but locally favored under drought. Strong genetic correlation between WUE and flowering time may facilitate the evolution of drought escape, or constrain independent evolution of these traits. Terminal drought favored drought escape in these spring flowering accessions of A. thaliana. WUE plasticity may be favored over completely fixed development in environments with periodic drought.