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Genotypic differences in transpiration rate responses to high vapour pressure deficit (VPD) was earlier reported. Here we tested the hypothesis that this limitation could relate to different degrees of dependence on the apoplastic (spaces between cells), and symplastic water transport pathways (through cells via aquaporin-facilitated transport), which are known to have different hydraulic conductivities. The low transpiration rate (Tr) genotype PRLT 2/89/33 either restricted its transpiration under high VPD, or was more sensitive to VPD than H77/833-2, when grown hydroponically or in soil. The slope of the transpiration response to an ascending series of VPD was lower in whole plants than in de-rooted shoots. In addition, the transpiration response of detached leaves to moderately high VPD (2.67kPa), normalised against leaves exposed to constant VPD (1.27kPa), was similar in low and high Tr genotypes. This suggested that roots hydraulics were a substantial limitation to water flow in pearl millet, especially under high VPD. The dependence on the apoplastic and symplastic water transport pathways was investigated by assessing the transpiration response of plants treated with inhibitors specific to the AQP-mediated symplastic pathway (AgNO3 and H2O2) and to the apoplastic pathway (precipitates of Cu(Fe(CN)6) or Cu(CuFe(CN)6)). When CuSO4 alone was used, Cu ions caused an inhibition of transpiration in both genotypes and more so in H77/833-2. The transpiration of high Tr H77/833-2 was decreased more by AQP inhibitors under low VPD (1.8kPa) than in PRLT 2/89/33, whereas under high VPD (4.2kPa), the transpiration of PRLT 2/89/33 was decreased more by AQP inhibitors than in H77/833-2. The transpiration rate of detached leaves from H77/833-2 when treated with AgNO3 decreased more than in PRLT 2/89/33. Although the root hydraulic conductivity of both genotypes was similar, it decreased more upon the application of a symplastic inhibitor in H77/833-2. The transpiration of low Tr PRLT 2/89/33 was decreased more by apoplastic inhibitors under both low and high VPD. Then the hydraulic conductivity decreased more upon the application of an apoplastic inhibitor in PRLT 2/89/33. In conclusion, both pathways contributed to water transport, and their contribution varied with environmental conditions and genotypes. Roots were a main source of hydraulic limitation in these genotypes of pearl millet, although a leaf limitation was not excluded. The similarity between genotypes in root hydraulic conductivity under normal conditions also suggests changes in this conductivity upon changes in the evaporative demand. The low Tr genotype depended more on the apoplastic pathway for water transport, whereas the high Tr genotype depended on both pathway, may be by 'tuning-up' the symplastic pathway under high transpiration demand, very likely via the involvement of aquaporins.

Chickpeas are often grown under receding soil moisture and suffer ~50% yield losses due to drought stress. The timing of soil water use is considered critical for the efficient use of water under drought and to reduce yield losses. Therefore the root growth and the soil water uptake of 12 chickpea genotypes known for contrasts in drought and rooting response were monitored throughout the growth period both under drought and optimal irrigation. Root distribution reduced in the surface and increased in the deep soil layers below 30cm in response to drought. Soil water uptake was the maximum at 45-60cm soil depth under drought whereas it was the maximum at shallower (15-30 and 30-45cm) soil depths when irrigated. The total water uptake under drought was 1-fold less than optimal irrigation. The amount of water left unused remained the same across watering regimes. All the drought sensitive chickpea genotypes were inferior in root distribution and soil water uptake but the timing of water uptake varied among drought tolerant genotypes. Superiority in water uptake in most stages and the total water use determined the best adaptation. The water use at 15-30cm soil depth ensured greater uptake from lower depths and the soil water use from 90-120cm soil was critical for best drought adaptation. Root length density and the soil water uptake across soil depths were closely associated except at the surface or the ultimate soil depths of root presence.

Traits influencing plant water use eventually define the fitness of genotypes for specific rainfall environments. We assessed the response of several water use traits to vapour pressure deficit (VPD) in pearl millet (Pennisetum glaucum (L.) R.Br.) genotypes known to differ in drought adaptation mechanisms: PRLT 2/89-33 (terminal drought-adapted parent), H 77/833-2 (terminal drought-sensitive parent) and four near-isogenic lines introgressed with a terminal drought tolerance quantitative trait locus (QTL) from PRLT 2/89-33 (ICMR01029, ICMR01031, ICMR02042, and ICMR02044). Plant water use traits at various levels of plant organisation were evaluated in seven experiments in plants exposed either transiently or over the long term to different VPD regimes: biomass components, transpiration (water usage per time unit) and transpiration rate (TR) upon transient VPD increase (gH2Ocm-2h-1)), transpiration efficiency (g dry biomass per kg H2O transpired), leaf expansion rate (cm per thermal time unit) and root anatomy (endodermis dimensions)). High VPD decreased biomass accumulation by reducing tillering, the leaf expansion rate and the duration of leaf expansion; decreased root endodermis cell size; and increased TR and the rate of TR increase upon gradual short-term VPD increases. Such changes may allow plants to increase their water transport capacity in a high VPD environment and are genotype-specific. Some variation in water use components was associated with terminal drought adaptation QTL. Knowledge of water use traits' plasticity in growth environments that varied in evaporative demand, and on their genetic determinacy, is necessary to develop trait-based breeding approaches to complex constraints.

The reproductive phase of chickpea (Cicer arietinum L.) is more sensitive to water deficits than the vegetative phase. The characteristics that confer drought tolerance to genotypes at the reproductive stage are not well understood; especially which characteristics are responsible for differences in seed yield under water stress. In two consecutive years, 10 genotypes with contrasting yields under terminal drought stress in the field were exposed to a gradual, but similar, water stress in the glasshouse. Flower number, flower+pod+seed abortion percentage, pod number, pod weight, seed number, seed yield, 100-seed weight (seed size), stem+leaf weight and harvest index (HI) were recorded in well watered plants (WW) and in water-stressed plants (WS) when the level of deficit was mild (phase I), and when the stress was severe (phase II). The WS treatment reduced seed yield, seed and pod number, but not flower+pod+seed abortion percentage or 100-seed weight. Although there were significant differences in total seed yield among the genotypes, the ranking of the seed yield in the glasshouse differed from the ranking in the field, indicating large genotype×environment interaction. Genetic variation for seed yield and seed yield components was observed in the WW treatment, which also showed differences across years, as well as in the WS treatment in both the years, so that the relative seed yield and relative yield components (ratio of values under WS to those under WW) were used as measures of drought tolerance. Relative total seed yield was positively associated with relative total flower number (R2=0.23 in year 2) and relative total seed number (R2=0.83, R2=0.79 in years 1 and 2 respectively). In phase I (mild stress), relative yield of seed produced in that phase was found to be associated with the flower number in both the years (R2=0.69, R2=0.76 respectively). Therefore, the controlled drought imposition that was used, where daily water loss from the soil was made equal for all plants, revealed genotypic differences in the sensitivity of the reproductive process to drought. Under these conditions, the seed yield differences in chickpea were largely related to the capacity to produce a large number of flowers and to set seeds, especially in the early phase of drought stress when the degree of water deficit was mild.

High temperatures and decreased rainfall are detrimental to yield in chickpea (Cicer arietinum L.), particularly during grain filling. This study aimed to (i) assess the individual and combined effects of drought and heat stress on biochemical seed-filling processes, (ii) determine genotypic differences in heat and drought tolerance, and (iii) determine any cross-tolerance. Plants were grown outdoors in the normal growing season when temperatures during seed filling were <32-20°C or were planted late (temperatures >32-20°C; heat stress). Half of the pots were kept adequately watered throughout, but water was withheld from the others from the initiation of seed filling until the relative leaf water content reached 50% of the irrigated plants (drought stress); all plants were rewatered thereafter until seed maturit. Water was withheld for 13 days (normal sowing) and 7 days (late sowing), so soil moisture decreased by 54-57%. Tests on leaves and seeds were performed after the stress. Individual and combined stress damaged membranes, and decreased cellular oxidising ability, stomatal conductance, PSII function and leaf chlorophyll content; damage was greater under combined stress. Leaf Rubisco activity increased with heat stress, decreased with drought stress and decreased severely with combined stress. Sucrose and starch concentrations decreased in all seeds through reductions in biosynthetic enzymes; reductions were greater under combined stress. These effects were more severe in heat- and drought-sensitive genotypes compared with drought-tolerant genotypes. Drought stress had a greater effect than heat stress on yield and the biochemical seed-filling mechanisms. Drought- and heat-tolerant genotypes showed partial cross-tolerance.

Under increasing water scarcity, food production for an increasing population is a global challenge. Maintaining crop production under limiting water supply is a common problem in agriculture, which is best addressed by the coordinated efforts of geneticists, physiologists and agronomists. This special issue is a selection of oral and poster presentations at the InterDrought IV conference, held in Perth (2-6 September 2013). These papers provide a broad, multidisciplinary view on the way to develop improved cultivars in the face of water deficit, providing the conference highlight: an integration of views from different disciplinary angles, generating constructive debate that was not buried in disciplinary silos. More specifically, the topics covered deal with the challenge of adaptation implicit in genotype-by-environment interaction, bring new perspectives on root systems and water productivity, and review the challenges and opportunities provided by crop management, genomic and transgenic approaches to cultivar improvement.

Post-rainy sorghum (Sorghum bicolor (L.) Moench) production underpins the livelihood of millions in the semiarid tropics, where the crop is affected by drought. Drought scenarios have been classified and quantified using crop simulation. In this report, variation in traits that hypothetically contribute to drought adaptation (plant growth dynamics, canopy and root water conducting capacity, drought stress responses) were virtually introgressed into the most common post-rainy sorghum genotype, and the influence of these traits on plant growth, development, and grain and stover yield were simulated across different scenarios. Limited transpiration rates under high vapour pressure deficit had the highest positive effect on production, especially combined with enhanced water extraction capacity at the root level. Variability in leaf development (smaller canopy size, later plant vigour or increased leaf appearance rate) also increased grain yield under severe drought, although it caused a stover yield trade-off under milder stress. Although the leaf development response to soil drying varied, this trait had only a modest benefit on crop production across all stress scenarios. Closer dissection of the model outputs showed that under water limitation, grain yield was largely determined by the amount of water availability after anthesis, and this relationship became closer with stress severity. All traits investigated increased water availability after anthesis and caused a delay in leaf senescence and led to a 'stay-green' phenotype. In conclusion, we showed that breeding success remained highly probabilistic; maximum resilience and economic benefits depended on drought frequency. Maximum potential could be explored by specific combinations of traits.

Terminal drought causes major yield loss in chickpea, so it is imperative to identify genotypes with best suited adaptive traits to secure yield in terminal drought-prone environments. Here, we evaluated chickpea (At) rd29A:: (At) DREB1A transgenic events (RD2, RD7, RD9 and RD10) and their untransformed C235 genotype for growth, water use and yield under terminal water-stress (WS) and well-watered (WW) conditions. The assessment was made across three lysimetric trials conducted in contained environments in the greenhouse (2009GH and 2010GH) and the field (2010F). Results from the greenhouse trials showed genotypic variation for harvest index (HI), yield, temporal pattern of flowering and seed filling, temporal pattern of water uptake across crop cycle, and transpiration efficiency (TE) under terminal WS conditions. The mechanisms underlying the yield gain in the WS transgenic events under 2009GH trial was related to conserving water for the reproductive stage in RD7, and setting seeds early in RD10. Water conservation also led to a lower percentage of flower and pod abortion in both RD7 and RD10. Similarly, in the 2010GH trial, reduced water extraction during vegetative stage in events RD2, RD7 and RD9 was critical for better seed filling in the pods produced from late flowers in RD2, and reduced percentage of flower and pod abortion in RD2 and RD9. However, in the 2010F trial, the increased seed yield and HI in RD9 compared with C235 came along only with small changes in water uptake and podding pattern, probably not causal. Events RD2 (2010GH), RD7 (2010GH) and RD10 (2009GH) with higher seed yield also had higher TE than C235. The results suggest that DREB1A, a transcription factor involved in the regulation of several genes of abiotic stress response cascade, influenced the pattern of water uptake and flowering across the crop cycle, leading to reduction in the percentage of flower and pod abortion in the glasshouse trials.

Water deficit is the main yield-limiting factor across the Asian and African semiarid tropics and a basic consideration when developing crop cultivars for water-limited conditions is to ensure that crop water demand matches season water supply. Conventional breeding has contributed to the development of varieties that are better adapted to water stress, such as early maturing cultivars that match water supply and demand and then escape terminal water stress. However, an optimisation of this match is possible. Also, further progress in breeding varieties that cope with water stress is hampered by the typically large genotype×environment interactions in most field studies. Therefore, a more comprehensive approach is required to revitalise the development of materials that are adapted to water stress. In the past two decades, transgenic and candidate gene approaches have been proposed for improving crop productivity under water stress, but have had limited real success. The major drawback of these approaches has been their failure to consider realistic water limitations and their link to yield when designing biotechnological experiments. Although the genes are many, the plant traits contributing to crop adaptation to water limitation are few and revolve around the critical need to match water supply and demand. We focus here on the genetic aspects of this, although we acknowledge that crop management options also have a role to play. These traits are related in part to increased, better or more conservative uses of soil water. However, the traits themselves are highly dynamic during crop development: they interact with each other and with the environment. Hence, success in breeding cultivars that are more resilient under water stress requires an understanding of plant traits affecting yield under water deficit as well as an understanding of their mutual and environmental interactions. Given that the phenotypic evaluation of germplasm/breeding material is limited by the number of locations and years of testing, crop simulation modelling then becomes a powerful tool for navigating the complexity of biological systems, for predicting the effects on yield and for determining the probability of success of specific traits or trait combinations across water stress scenarios.

Terminal drought is a major constraint to chickpea productivity. Carbon isotope discrimination (Δ13C), an integrator of plant behaviour influencing transpiration efficiency (TE), is an important component of yield under drought. The variation in Δ13C and its association with yield was assessed in the reference collection of chickpea germplasm. Drought stress reduced shoot biomass by 36-39% and grain yield by 23%. Mean Δ13C was low and the range of genetic variation was high under drought stress. Largely, high Δ13C accessions were early in flowering (40-50 days), moderate in shoot biomass, high in seed yields and high in harvest index (HI). Δ13C was positively correlated with seed yield in both the years under drought stress, only in 2008-09 under optimal irrigation. This positive association was very close with HI. Among the yield components, Δ13C was closely associated with pod numbers per unit area and seed size under drought stress. Path coefficients showed no direct association of Δ13C with grain yield but an indirect negative association through shoot biomass at maturity and a close positive association through HI. The closest association of HI or shoot biomass was seen in the maturity group of accessions that experienced the optimum terminal drought stress.

In addition to characterising root architecture, evaluating root water uptake ability is important for understanding drought response. A series of three lysimeter studies were conducted using the OryzaSNP panel, which consists of 20 diverse rice (Oryza sativa L.) genotypes. Large genotypic differences in drought response were observed in this genotype panel in terms of plant growth and water uptake. Total water uptake and daily water uptake rates in the drought-stress treatment were correlated with root length density, especially at depths below 30cm. Patterns of water uptake among genotypes remained consistent throughout the stress treatments: genotypes that initially extracted more water were the same genotypes that extracted more water at the end of the study. These results suggest that response to drought by deep root growth, rather than a conservative soil water pattern, seems to be important for lowland rice. Genotypes in the O. sativa type aus group showed some of the greatest water uptake and root growth values. Since the OryzaSNP panel has been genotyped in detail with SNP markers, we expect that these results will be useful for understanding the genetics of rice root growth and function for water uptake in response to drought.

As water availability is critical for reproduction, terminal drought tolerance may involve water-saving traits. Experiments were undertaken under different vapour pressure deficit (VPD) and water regimes (water stress (WS) and well watered (WW)) to test genotypic differences and trait relationships in the fraction of transpirable soil water (FTSW) at which transpiration declines, canopy conductance (proxied by transpiration rate (TR, g H2Ocm-2h-1)), canopy temperature depression (CTD, °C), transpiration efficiency (TE, gkg-1) and growth parameters, using 15 contrasting cowpea (Vigna unguiculata (L.) Walp.) genotypes. Under WW conditions at the vegetative and early podding stages, plant mass and leaf area were larger under low VPD, and was generally lower in tolerant than in sensitive genotypes. Several tolerant lines had lower TR under WW conditions and restricted TR more than sensitive lines under high VPD. Under WS conditions, transpiration declined at a lower FTSW in tolerant than in sensitive lines. Tolerant lines also maintained higher TR and CTD under severe stress. TE was higher in tolerant genotypes under WS conditions. Significant relationships were found between TR, and TE, CTD and FTSW under different water regimes. In summary, traits that condition how genotypes manage limited water resources discriminated between tolerant and sensitive lines. Arguably, a lower canopy conductance limits plant growth and plant water use, and allows tolerant lines to behave like unstressed plants until the soil is drier and to maintain a higher TR under severe stress, as lower TR at high VPD leads to higher TE.

Pearl millet (Pennisetum glaucum (L.) R.Br.) is a resilient crop suiting the harshest conditions of the semi-arid tropics, in which we assessed possible relationships between crop tolerance, anti-oxidative enzyme activity and plant/soil water status. Biochemical acclimation and cell homeostasis traits have been proposed as critical for the drought tolerance of crops, but their limited practical application in breeding so far suggests that the role of biochemical acclimation for drought tolerance is still unclear. Previous research may have been of limited value because it has not approached biochemical acclimation from the angle of plant water relations. Four pearl millet genotypes, contrasting for terminal drought tolerance, were evaluated (sensitive H77/833-2, tolerant PRLT2/89-33 and two near isogenic lines carrying a terminal drought tolerance quantitative trait locus) under water-stress (WS) and well-watered (WW) conditions in a lysimetric system that simulates field-like conditions. We assessed the genotypic variation and relationship between photosynthetic pigments (chlorophylls a and b and carotenoids), antioxidative isoenzymatic spectrum (superoxide dismutase, ascorbate peroxidase and catalase), physiological traits (soil moisture available, normalised transpiration, stay-green score and water extraction) and biomass and yield. Biochemical traits investigated were tightly related among each other under WS conditions but not under WW conditions. Two major ascorbate peroxidase isoforms (APX6&7), whose variation in both water regimes reflected the presence/absence of the drought tolerance quantitative trait locus, were identified, but these did not relate to yield. Both, yield and biochemical traits under terminal drought stress were closely related to the traits linked to plant/soil water status (soil moisture available, normalised transpiration, stay-green score and water extraction), whereas yield and the biochemical indicators were not correlated, except for one. It is concluded that there is no direct effect of biochemical traits on yield parameters since both are consequences of soil-plant water status and their putative relation appear to be secondary - through plant/soil water status.

Intermittent drought, which varies in intensity, severely limits groundnut (Arachis hypogaea L.) yields. Experiments were conducted to assess root development, water uptake, transpiration efficiency, yield components and their relationships, in 20 groundnut genotypes under well watered (WW), and mild (DS-1), medium (DS-2) and severe (DS-3) intermittent stress. Pod yield decreased 70%, 55% and 35% under severe, medium and mild stress, respectively. Pod yield varied among genotypes, and showed significant genotype-by-treatment effects. Root length density (RLD) varied among genotypes before and after stress, although RLD did not discriminate tolerant from sensitive lines. Total water uptake and RLD under water stress had a weakly significant relationship. Water extraction from the soil profile was highest under severe stress. Water uptake varied among genotypes in all water regimes, but correlated with pod yield under WW conditions. The relative harvest index (HI) (i.e. the ratio of the HI under stress to HI under WW conditions) was closely related to the pod yield in all three intermittent stresses (R2=0.68 in DS-1; R2=0.65 in DS-2; R2=0.86 in DS-3) and was used as an index of stress tolerance. Under medium and severe stresses, the relative HI was negatively related to plant leaf weight (R2=0.79 in DS-2; R2=0.53 in DS-3), but less so under mild stress (R2=0.31). The results suggest that under intermittent stress, genotypes with a lower leaf area may use water more sparingly during the drying cycle with less damaging consequences for reproduction and pod.

A stay-green phenotype enhances the adaptation of sorghum (Sorghum bicolor (L.) Moench) to terminal drought, although the mechanisms leading to its expression remain unclear. Differences in tillering and leaf area at anthesis, transpiration efficiency (TE), water extraction, harvest index (HI) and yield under terminal drought and fully irrigated conditions were assessed in 29 introgression lines (IL) targeting stay-green quantitative trait loci (QTLs) Stg1, Stg2, Stg3, Stg4, StgA and StgB in an S35 background, and 16 IL targeting Stg1, Stg3, Stg4 and StgB in an R16 background. TE was increased by StgB in the R16 background, whereas there was no effect in the S35 background. Water extraction was increased by Stg1 in the S35 background but not in R16. StgB modified the proportion of water extracted before and after anthesis in the S35 background. While tillering and leaf area at anthesis were decreased by Stg1 and Stg3 in S35, there was no effect in R16. Yield data under fully irrigated conditions showed higher tiller grain yield in Stg1, Stg2 and Stg3 ILs. Although yield differences were mostly explained by HI variation, the yield variation unexplained by HI was closely related to TE in S35 (R2=0.29) and R16 (R2=0.72), and was closely related to total water extracted in S35 (R2=0.41) but not in R16. These data indicate the potential for several stay-green QTLs to affect traits related to plant water use. However, these effects depend on the interaction between the genetic background and individual QTLs.

Chickpea (Cicer arietinum L.) is often exposed to terminal drought, and deep, profuse rooting has been proposed as the main breeding target to improve terminal drought tolerance. This work tested whether plant water use at the vegetative stage and under non-limited water conditions could relate to the degree of sensitivity of chickpea to terminal drought. Transpiration response to a range of vapour pressure deficits under controlled and outdoor conditions was measured with canopy conductance using gravimetric measurements and thermal imagery in eight chickpea genotypes with comparable phenology and contrasting seed yield under terminal drought in the field. Additionally, the response of plant growth and transpiration to progressive soil moisture depletion was assayed in the same genotypes. Drought-tolerant genotypes had a lower canopy conductance under fully irrigated conditions at the vegetative stage; this trend was reversed at the early pod filling stage. While two sensitive genotypes had high early growth vigour and leaf development, there was a trend of lower growth in tolerant genotypes under progressive soil drying than in sensitive ones. Tolerant genotypes also exhibited a decline of transpiration in wetter soil compared to sensitive genotypes. Canopy conductance could be proxied by measuring leaf temperature with an infrared camera, although the relationship lost sensitivity at the pod filling stage. This work suggests that some traits contribute to water saving when water does not limit plant growth and development in drought-tolerant chickpea. It is hypothesised that this water would be available for the reproduction and grain filling stages.