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Warming nights are correlated with declining wheat growth and yield. As a key determinant of plant biomass, respiration consumes O2 as it produces ATP and releases CO2 and is typically reduced under warming to maintain metabolic efficiency. We compared the response of respiratory O2 and CO2 flux to multiple night and day warming treatments in wheat leaves and roots, using one commercial (Mace) and one breeding cultivar grown in controlled environments. We also examined the effect of night warming and a day heatwave on the capacity of the ATP-uncoupled alternative oxidase (AOX) pathway. Under warm nights, plant biomass fell, respiratory CO2 release measured at a common temperature was unchanged (indicating higher rates of CO2 release at prevailing growth temperature), respiratory O2 consumption at a common temperature declined, and AOX pathway capacity increased. The uncoupling of CO2 and O2 exchange and enhanced AOX pathway capacity suggest a reduction in plant energy demand under warm nights (lower O2 consumption), alongside higher rates of CO2 release under prevailing growth temperature (due to a lack of down-regulation of respiratory CO2 release). Less efficient ATP synthesis, teamed with sustained CO2 flux, could thus be driving observed biomass declines under warm nights.

QTL controlling vigour and related traits were identified in a chickpea RIL population and validated in diverse sets of germplasm. Robust KASP markers were developed for marker-assisted selection. To understand the genetic constitution of vigour in chickpea (Cicer arietinum L.), genomic data from a bi-parental population and multiple diversity panels were used to identify QTL, sequence-level haplotypes and genetic markers associated with vigour-related traits in Australian environments. Using 182 Recombinant Inbred Lines (RILs) derived from a cross between two desi varieties, Rupali and Genesis836, vigour QTL independent of flowering time were identified on chromosomes (Ca) 1, 3 and 4 with genotypic variance explained (GVE) ranging from 7.1 to 28.8%. Haplotype analysis, association analysis and graphical genotyping of whole-genome re-sequencing data of two diversity panels consisting of Australian and Indian genotypes and an ICRISAT Chickpea Reference Set revealed a deletion in the FTa1–FTa2–FTc gene cluster of Ca3 significantly associated with vigour and flowering time. Across the RIL population and diversity panels, the impact of the deletion was consistent for vigour but not flowering time. Vigour-related QTL on Ca4 co-located with a QTL for seed size in Rupali/Genesis836 (GVE = 61.3%). Using SNPs from this region, we developed and validated gene-based KASP markers across different panels. Two markers were developed for a gene on Ca1, myo -inositol monophosphatase (CaIMP), previously proposed to control seed size, seed germination and seedling growth in chickpea. While associated with vigour in the diversity panels, neither the markers nor broader haplotype linked to CaIMP was polymorphic in Rupali/Genesis836. Importantly, vigour appears to be controlled by different sets of QTL across time and with components which are independent from phenology.

High temperature stress inhibits photosynthesis and threatens wheat production. One measure of photosynthetic heat tolerance is Tcrit – the critical temperature at which incipient damage to photosystem II (PSII) occurs. This trait could be improved in wheat by exploiting genetic variation and genotype-by-environment interactions (GEI). Flag leaf Tcrit of 54 wheat genotypes was evaluated in 12 thermal environments over 3 years in Australia, and analysed using linear mixed models to assess GEI effects. Nine of the 12 environments had significant genetic effects and highly variable broad-sense heritability (H2 ranged from 0.15 to 0.75). Tcrit GEI was variable, with 55.6% of the genetic variance across environments accounted for by the factor analytic model. Mean daily growth temperature in the month preceding anthesis was the most influential environmental driver of Tcrit GEI, suggesting biochemical, physiological and structural adjustments to temperature requiring different durations to manifest. These changes help protect or repair PSII upon exposure to heat stress, and may improve carbon assimilation under high temperature. To support breeding efforts to improve wheat performance under high temperature, we identified genotypes superior to commercial cultivars commonly grown by farmers, and demonstrated potential for developing genotypes with greater photosynthetic heat tolerance.

Genotypic variation in transpiration (Tr) response to vapour pressure deficit (VPD) has been studied in many crop species. There is debate over whether shoots or roots drive these responses. We investigated how stomata coordinate with plant hydraulics to mediate Tr response to VPD and influence leaf water status in wheat (Triticum aestivum L.). We measured Tr and stomatal conductance (gs) responses to VPD in well-watered, water-stressed and de-rooted shoots of eight wheat genotypes. Tr response to VPD was related to stomatal sensitivity to VPD and proportional to gs at low VPD, except in the water-stressed treatment, which induced strong stomatal closure at all VPD levels. Moreover, gs response to VPD was driven by adaxial stomata. A simple linear Tr response to VPD was associated with unresponsive gs to VPD. In contrast, segmented linear Tr to VPD response was mostly a function of gs with the breakpoint depending on the capacity to meet transpirational demand and set by the shoots. However, the magnitude of Tr response to VPD was influenced by roots, soil water content and stomatal sensitivity to VPD. These findings, along with a theoretical model suggest that stomata coordinate with plant hydraulics to regulate Tr response to VPD in wheat.

Heat-induced inhibition of photosynthesis is a key factor in declining wheat performance and yield. Variation in wheat heat tolerance can be characterised using the critical temperature (Tcrit) above which incipient damage to the photosynthetic machinery occurs. We investigated intraspecies variation and plasticity of wheat Tcrit under elevated temperature in field and controlled environment experiments. We also assessed whether intraspecies variation in wheat Tcrit mirrors patterns of global interspecies variation in heat tolerance reported for mostly wild, woody plants. In the field, wheat Tcrit varied through the course of a day, peaking at noon and lowest at sunrise, and increased as plants developed from heading to anthesis and grain filling. Under controlled temperature conditions, heat stress (36°C) was associated with a rapid rise in wheat Tcrit (i.e. within two hours of heat stress) that peaked after 3–4 days. These peaks in Tcrit indicate a physiological limitation to photosystem II heat tolerance. Analysis of a global dataset (comprising 183 Triticum and wild wheat (Aegilops) species) generated from the current study and a systematic literature review showed that wheat leaf Tcrit varied by up to 20°C (about two-thirds of reported global plant interspecies variation). However, unlike global patterns of interspecies Tcrit variation which has been linked to latitude of genotype origin, intraspecific variation in wheat Tcrit was unrelated to that. Yet, the observed genotypic variation and plasticity of wheat Tcrit suggests that this trait could be a useful tool for high-throughput phenotyping of wheat photosynthetic heat tolerance.

QTL controlling vigour and related traits were identified in a chickpea RIL population and validated in diverse sets of germplasm. Robust KASP markers were developed for marker-assisted selection. To understand the genetic constitution of vigour in chickpea (Cicer arietinum L.), genomic data from a bi-parental population and multiple diversity panels were used to identify QTL, sequence-level haplotypes and genetic markers associated with vigour-related traits in Australian environments. Using 182 Recombinant Inbred Lines (RILs) derived from a cross between two desi varieties, Rupali and Genesis836, vigour QTL independent of flowering time were identified on chromosomes (Ca) 1, 3 and 4 with genotypic variance explained (GVE) ranging from 7.1 to 28.8%. Haplotype analysis, association analysis and graphical genotyping of whole-genome re-sequencing data of two diversity panels consisting of Australian and Indian genotypes and an ICRISAT Chickpea Reference Set revealed a deletion in the FTa1-FTa2-FTc gene cluster of Ca3 significantly associated with vigour and flowering time. Across the RIL population and diversity panels, the impact of the deletion was consistent for vigour but not flowering time. Vigour-related QTL on Ca4 co-located with a QTL for seed size in Rupali/Genesis836 (GVE = 61.3%). Using SNPs from this region, we developed and validated gene-based KASP markers across different panels. Two markers were developed for a gene on Ca1, myo -inositol monophosphatase (CaIMP), previously proposed to control seed size, seed germination and seedling growth in chickpea. While associated with vigour in the diversity panels, neither the markers nor broader haplotype linked to CaIMP was polymorphic in Rupali/Genesis836. Importantly, vigour appears to be controlled by different sets of QTL across time and with components which are independent from phenology.

Climate change and future warming will significantly affect crop yield. The capacity of crops to dynamically adjust physiological processes (i.e. acclimate) to warming might improve overall performance. Understanding and quantifying the degree of acclimation in field crops could ensure better parameterization of crop and Earth System models and predictions of crop performance. We hypothesized that for field-grown wheat, when measured at a common temperature (25°C), crops grown under warmer conditions would exhibit acclimation, leading to enhanced crop performance and yield. Acclimation was defined as: (i) decreased rates of net photosynthesis at 25°C (A25) coupled with lower maximum carboxylation capacity (Vcmax25); (ii) reduced leaf dark respiration at 25°C (both in terms of O2 consumption, Rdark_O225; and CO2 efflux, Rdark_CO225); and (iii) lower Rdark_CO225:Vcmax25. Field experiments were conducted over two seasons with 20 wheat genotypes, sown at three different planting dates, to test these hypotheses. Leaf-level CO2 based traits (A25, Rdark_CO225, and Vcmax25) did not show the classic acclimation responses that we hypothesized; by contrast, the hypothesized changes in Rdark_O2 were observed. These findings have implications for predictive crop models that assume similar temperature response among these physiological processes, and for predictions of crop performance in a future warmer world.

Plants are inherently dynamic. Dynamics minimise stress while flexibly acquiring resouces. Three examples are presented for plants tolerating saline soil: transport of NaCl, water and macronutrients is non‐uniform along a branched root, visualised by mapping expressions with tissues and age; water and NaCl redistribute between shoot and soil at nighttime; and ATP for salt exclusion is much lower in thinner branch than main roots, quantified using a biophysical model and geometry from anatomy. Non‐invasive phenotyping and precision agriculture technologies can be used together to harness dynamics, but analytical methods are needed. A plant advancing in time through a soil and atmosphere space is proposed as a framework for dynamic data and their relationship to crop improvement. Refereed/Peer-reviewed

Emmer wheat (Triticum dicoccon Schrank) is a potential source of new genetic diversity for the improvement of hexaploid bread wheat. Emmer wheat was crossed and backcrossed to bread wheat and 480 doubled haploids (DHs) were produced from BC1F1 plants with hexaploid appearance derived from 19 crossses. These DHs were screened under well-watered conditions (E1) in 2013 to identify high-yielding materials with similar phenology. One-hundred and eighty seven DH lines selected on this basis, 4 commercial bread wheat cultivars and 9 bread wheat parents were then evaluated in extensive field experiments under two contrasting moisture regimes in north-western NSW in 2014 and 2015. A significant range in the water-use-efficiency of grain production (WUEGrain) was observed among the emmer derivatives. Of these, 8 hexaploid lines developed from 8 different emmer wheat parents had significantly improved intrinsic water-use-efficiency (WUEintr) and instantaneous water-use-efficiency (WUEi) compared to their bread wheat recurrent parents. Accurate and large scale field-based phenotyping was effective in identifying emmer wheat derived lines with superior performance to their hexaploid bread wheat recurrent parents under moisture stress.

High temperatures account for major wheat yield losses annually and, as the climate continues to warm, these losses will probably increase. Both photosynthesis and respiration are the main determinants of carbon balance and growth in wheat, and both are sensitive to high temperature. Wheat is able to acclimate photosynthesis and respiration to high temperature, and thus reduce the negative affects on growth. The capacity to adjust these processes to better suit warmer conditions stands as a potential avenue toward reducing heat-induced yield losses in the future. However, much remains to be learnt about such phenomena. Here, we review what is known of high temperature tolerance in wheat, focusing predominantly on the high temperature responses of photosynthesis and respiration. We also identify the many unknowns that surround this area, particularly with respect to the high temperature response of wheat respiration and the consequences of this for growth and yield. It is concluded that further investigation into the response of photosynthesis and respiration to high temperature could present several methods of improving wheat high temperature tolerance. Extending our knowledge in this area could also lead to more immediate benefits, such as the enhancement of current crop models.

Wheat is sensitive to heat stress, particularly during grain filling, and this reduces grain yield. Ancestral wheat species, such as emmer wheat (Triticum dicoccon Schrank), represent potential sources of new genetic diversity for traits that may impact wheat responses to heat stress. However, the diversity available in emmer wheat has only been explored superficially. Recently developed emmer derived hexaploid wheat genotypes were evaluated for physiological, phenological and agronomic traits in a multi-environment, multi-season strategy. The emmer-based hexaploid lines were developed from crosses and backcrosses to 9 hexaploid recurrent parents and these genotypes and 7 commercial cultivars were evaluated under two times of sowing (E1 and E2) in the field for three consecutive years (2014-2016). The materials were genotyped using a 90 K SNP platform and these data used to estimate the contribution of emmer wheat to the progeny. Significant phenotypic and genetic variation for traits were observed. Higher temperature during reproductive development and grain filling reduced trait expression. Emmer progeny with greater trait values than their recurrent parents and commercial cultivars in both environments were found. Derivatives with higher physiological trait values yielded well in both environments; as indicated by the clustering of genotypes. The emmer wheat parent contributed between 1 and 43 % of the genome of the emmer-based hexaploid progeny, and progeny with greater emmer contribution had superior trait values in both environments. These results showed a positive effect of direct emmer introgression on wheat performance under heat stress. Mitigation of high temperature stress through the introgression of favorable alleles from wheat close relatives into modern wheat cultivars is possible.

This study investigated the relationship between osmotic potential at full hydration (π100) and turgor loss point (ΨTLP) in wheat (Triticum aestivum) to determine the potential of using π100 to predict ΨTLP under well-watered (WW) and drought (WS) conditions. Two methods for determining π100 were tested: pressure–volume (PV) analysis and freezing point osmometry. The study also measured π100 in a range of 38 field-grown wheat cultivars to determine whether there is genetic variation in π100 under field conditions. π100 correlated with ΨTLP using both methods under both water treatments, particularly WS. Genetic variation of π100 in the field, under rainfed conditions, was greater than controlled conditions and ranged from −0.94 to −1.95 MPa. Overall, the evidence supports development of π100 as a novel tool for plant breeders to screen large populations of wheat and identify genotypes with lower ΨTLP, an integrative trait that is related to drought tolerance.

High temperature has a negative impact on wheat grain quality and reduces market value. Emmer wheat (Triticum dicoccon Schrank), one of the earliest domesticated wheat species, is a source of genetic diversity for the improvement of heat and drought tolerance in modern wheat. However, the potential of emmer wheat for the improvement of grain physical quality under high temperature stress is little studied. A diverse set of 184 emmer-based hexaploid lines was developed by crossing emmer wheat with hexaploid wheat and backcrossing once to hexaploid wheat. These materials, seven hexaploid recurrent parents and seven commercial cultivars, were evaluated at two times of sowing (E1 and E2) in the field, in 2015–2016. The materials were genotyped using a 90 K SNP platform and these data were used to estimate the contribution of emmer wheat to the progeny. Significant phenotypic and genetic variation for grain physical quality traits including protein content and test weight was observed. High temperature significantly increased protein content and decreased test weight. Large scale field phenotyping identified emmer progenies with improved grain characteristic compared to their respective parents and commercial cultivars in both environments. A few families consistently produced higher trait means across environments compared to their recurrent parents. The emmer wheat parent contributed between 1 and 37% of the genome in emmer-based genotypes. Selected emmer derived lines with superior protein content and test weight, tended to have a greater genetic contribution from the emmer parent, ranging from 12 to 37% and 7–37% in E1 and E2, respectively. It was concluded that new genetic variation for seed traits, such as protein content and test weight, can be introduced to hexaploid wheat from emmer wheat. The newly developed emmer derivatives identified with enhanced grain quality under high temperature stress can potentially be used to improve grain quality through breeding.

The response of wheat to the variables of climate change includes elevated CO2, high temperature, and drought which vary according to the levels of each variable and genotype. Independently, elevated CO2, high temperature, and terminal drought affect wheat biomass and grain yield, but the interactive effects of these three variables are not well known. The aim of this study was to determine the effects of elevated CO2 when combined with high temperature and terminal drought on the high-yielding traits of restricted-tillering and vigorous growth. It was hypothesized that elevated CO2 alone, rather than combined with high temperature, ameliorates the effects of terminal drought on wheat biomass and grain yield. It was also hypothesized that wheat genotypes with more sink capacity (e.g. high-tillering capacity and leaf area) have more grain yield under combined elevated CO2, high temperature, and terminal drought. Two pairs of sister lines with contrasting tillering and vigorous growth were grown in poly-tunnels in a four-factor completely randomized split-plot design with elevated CO2 (700 µL L(-1)), high day time temperature (3 °C above ambient), and drought (induced from anthesis) in all combinations to test whether elevated CO2 ameliorates the effects of high temperature and terminal drought on biomass accumulation and grain yield. For biomass and grain yield, only main effects for climate change variables were significant. Elevated CO2 significantly increased grain yield by 24-35% in all four lines and terminal drought significantly reduced grain yield by 16-17% in all four lines, while high temperature (3 °C above the ambient) had no significant effect. A trade-off between yield components limited grain yield in lines with greater sink capacity (free-tillering lines). This response suggests that any positive response to predicted changes in climate will not overcome the limitations imposed by the trade-off in yield components.

Yields of grain legumes are constrained by available water. Thus, it is crucial to understand traits influencing water uptake and the efficiency of using water to produce biomass. Global comparisons and comparisons at specific locations reveal that water use of different grain legumes is very similar, which indicates that water use efficiency varies over a wide range due to differences in biomass and yield. Moreover, yield increases more per millimetre of water used in cool season grain legumes than warm season species. Although greater contrasts have been observed across species and genotypes at the pot and lysimeter level, agronomic factors need to be taken into account when scaling those studies to field-level responses. Conservative water use strategies in grain legumes such as low stomatal conductance as approximated by low photosynthetic carbon isotope discrimination reduces yield potential, whereas temporal adjustments of stomatal conductance within the growing season and in response to environmental factors (such as vapour pressure deficit) helps to optimize the trade-off between carbon gain and water loss. Furthermore, improved photosynthetic capacity, reduced mesophyll conductance, reduced boundary layer, and re-fixation of respired CO2 were identified as traits that are beneficial without water deficit, but also under terminal and transient drought. Genotypic variability in some grain legume species has been observed for several traits that influence water use, water use efficiency, and yield, including root length and the temporal pattern of water use, but even more variation is expected from wild relatives. Albeit that N2 fixation decreases under drought, its impact on water use is still largely unknown, but the nitrogen source influences gas exchange and, thus, transpiration efficiency. This review concludes that conservative traits are needed under conditions of terminal drought to help maintain soil moisture until the pod-filling period, but profligate traits, if tightly regulated, are important under conditions of transient drought in order to profit from short intermittent periods of available soil moisture.

Increases in atmospheric carbon dioxide concentration ([CO2]) and temperature associated with future climates are expected to affect wheat growth and grain yield. The ability of wheat to adapt to these changes is close related to the response of the root system. The effect of elevated CO2, high temperature and the interaction elevated CO2 × high temperature (3 °C above the ambient temperature) on the growth and proliferation of the root system of two spring wheat genotypes differing in vigorous growth was evaluated. The breeding line Vigor 18 selected for vigorous shoot and root growth and the commercial cultivar Janz were grown in rhizo-boxes inside specially designed tunnel houses under ambient (390–400 μL L−1) and elevated CO2 (700 μL L−1), and ambient and 3 °C above the ambient temperature. Growth and proliferation of the root system was monitored through root mapping. Elevated CO2 enhanced root and shoot biomass, but this positive effect was reduced when plants were grown under high temperature. Total root length was also increased by elevated CO2, but only when plants were grown under ambient rather than under high temperature. Root growth response to elevated CO2 differed between genotypes; elevated CO2 stimulated root growth in the non-vigorous cultivar Janz, but not in vigorous liner Vigor 18. High temperature reduces the positive effect that elevated CO2 has on root growth and proliferation.

Wheat grown in the Mediterranean-type environments of southern Australia is often exposed to end-of-season drought (terminal drought). During the development of terminal drought, soil dries from the top of the profile exposing the upper part of the root system to water stress while deeper roots may still be able to access deeper soil water for grain filling. It is hypothesised that the part of the root system exposed to drying soil signals abscisic acid (ABA) production and the corresponding rise in ABA concentration in leaves causes partial stomatal closure, regulating the extraction of available water at depth. In the first step to test this hypothesis, a glasshouse experiment was conducted to identify contrasting stomatal response to terminal drought and production of ABA using four wheat genotypes adapted to different soil moisture environments. Terminal drought was induced by withholding water from anthesis in one half (WD) or both halves (DD) of the root system using split pots. Stomatal conductance decreased in all four genotypes in WD plants, but leaf ABA concentration and leaf water status differed. The cultivar Drysdale had higher leaf ABA concentrations and lower stomatal conductance, but leaf water status decreased. Leaf ABA concentration did not increase in WD plants of the breeding line IGW-3262, but stomatal conductance decreased and leaf water status was unchanged. All the genotypes behaved similarly under DD conditions, with increased leaf ABA concentration, lower stomatal conductance and severely dehydrated leaves. The possible causes of the above differences in leaf ABA concentration despite similar stomatal behaviour in wheat genotypes are discussed.

Understanding crop responses to increasing atmospheric CO2 requires knowledge of how their root systems grow, proliferate and function. The effect of elevated CO2 on the growth and proliferation of wheat root system (Triticum aestivum L.), was examined. Two pairs of sister lines of wheat contrasting in vigour (CV97 and CV207) and tillering (7750N and 7750PF) were grown in rhizo-boxes under ambient (380 μl L−1) and elevated CO2 (700 μl L−1), and the root growth and proliferation mapped. Elevated CO2 effects on shoot and root biomass were observed in the lines contrasting for vigour, but not in the lines contrasting for tillering. Root biomass was reduced by 67 % in the high vigour line CV97, reducing total plant biomass by 26 % compared to the low vigour line, CV207. This was due to a reduction in root length down the 1 m soil profile and root proliferation in the top 0.2 m layer. The reduction in root biomass was not compensated by an increase in shoot biomass. The reduction in root biomass under elevated CO2 in the vigour line CV97 can be explained through its inability to increase the sink strength due to the failure to increase tiller number to which the plant presumably responded by increasing losses of the newly assimilated carbon by respiration.

Background and aims Being able to monitor the hydration status of a plant would be useful to breeding programs and to providing insight into adaptation to water-limited environments, but most current methods are destructive or laborious. We evaluated novel non-invasive pressure probes (commercial name: ZIM-probe) for their potential in monitoring the water status of wheat (Triticum aestivum L.) leaves.

The United Nations declared 2016 as the International Year of Pulses (grain legumes) under the banner ‘nutritious seeds for a sustainable future’. A second green revolution is required to ensure food and nutritional security in the face of global climate change. Grain legumes provide an unparalleled solution to this problem because of their inherent capacity for symbiotic atmospheric nitrogen fixation, which provides economically sustainable advantages for farming. In addition, a legume-rich diet has health benefits for humans and livestock alike. However, grain legumes form only a minor part of most current human diets and legume crops are greatly under-utilized. Food security and soil fertility could be significantly improved by greater grain legume usage and increased improvement of a range of grain legumes. The current lack of coordinated focus on grain legumes has compromised human health, nutritional security and sustainable food production.

Post-anthesis water use is important for grain yield in wheat under drought because this water is immediately used for grain filling. The aim of this study was to determine whether root capacity for water uptake from deeper layers in the soil profile differed between two genotypes with contrasting stomatal behaviour under terminal drought. The wheat cultivar Drysdale and the breeding line IGW-3262 were grown in 1 m deep pots in a glasshouse under well-watered conditions until anthesis, when three watering treatments were imposed: (i) watering maintained at 90% pot soil water capacity (WW), (ii) watering withheld but supplementary watering supplied to the bottom 30 cm of the pot to keep this layer of the soil profile wet until physiological maturity (WB) and (iii) watering completely withheld (WS). Stomatal conductance, post-anthesis water use and water use efficiency, and grain yield were measured. Post-anthesis water use in Drysdale was similar in the WB and WW treatments, while in IGW-3262 it was 30% less in the WB treatment than in the WW treatment. In the WB treatment as the top soil dried, stomatal closure was faster in IGW-3262 than in Drysdale, which may have affected the capacity of roots to uptake available water at depth. The reduction in post-anthesis water use in IGW-3262 resulted in a decline in grain yield.

Ambient temperatures have increased since the beginning of the century and are predicted to continue rising under climate change. Such increases in temperature can cause heat stress: a severe threat to wheat production in many countries, particularly when it occurs during reproductive and grain-filling phases. Heat stress reduces plant photosynthetic capacity through metabolic limitations and oxidative damage to chloroplasts, with concomitant reductions in dry matter accumulation and grain yield. Genotypes expressing heat shock proteins are better able to withstand heat stress as they protect proteins from heat-induced damage. Heat tolerance can be improved by selecting and developing wheat genotypes with heat resistance. Wheat pre-breeding and breeding may be based on secondary traits like membrane stability, photosynthetic rate and grain weight under heat stress. Nonetheless, improvement in grain yield under heat stress implies selecting genotypes for grain size and rate of grain filling. Integrating ph...

Soil water (θ) dynamics are important parameters to monitor in any field-based drought research. Although apparent electrical conductivity (ECa) measured by electromagnetic (EM) induction has been used to estimate θ, little research has shown its successful application at the plot-scale for evaluating crop water use. An EM38 conductivity meter was used to collect time-lapse ECa data at the plot scale across a field cropped with 36 different chickpea genotypes. An empirical multiple linear regression model was established to predict θ measured by neutron probes and depth-specific electrical conductivity (σ) generated by a 1-D EM inversion algorithm. Soil water dynamics and movement were successfully mapped with a coefficient of determination (R2) of 0.87 and root-mean-square-error of 0.037 m3 m−3. The rate of soil drying varied with depth and was influenced by chickpea growth stages and genotypes. The results were also used to evaluate the differences in soil water use and rooting depths within- and across-plant species and during the growth stages. Coupled with physiology measurements, the approach can also be used to identify mechanisms of drought tolerance in the field and screening for effective water use in crop breeding programs.

The contrasting hydraulic properties of wheat (Triticum aestivum), narrow-leafed lupin (Lupinus angustifolius), and yellow lupin (Lupinus luteus) roots were identified by integrating measurements of water flow across different structural levels of organization with anatomy and modeling. Anatomy played a major role in root hydraulics, influencing axial conductance (L ax) and the distribution of water uptake along the root, with a more localized role for aquaporins (AQPs). Lupin roots had greater L ax than wheat roots, due to greater xylem development. L ax and root hydraulic conductance (L r) were related to each other, such that both variables increased with distance from the root tip in lupin roots. L ax and L r were constant with distance from the tip in wheat roots. Despite these contrasting behaviors, the hydraulic conductivity of root cells (Lp c) was similar for all species and increased from the root surface toward the endodermis. Lp c was largely controlled by AQPs, as demonstrated by dramatic reductions in Lp c by the AQP blocker mercury. Modeling the root as a series of concentric, cylindrical membranes, and the inhibition of AQP activity at the root level, indicated that water flow in lupin roots occurred primarily through the apoplast, without crossing membranes and without the involvement of AQPs. In contrast, water flow across wheat roots crossed mercury-sensitive AQPs in the endodermis, which significantly influenced L r. This study demonstrates the importance of examining root morphology and anatomy in assessing the role of AQPs in root hydraulics.

Hydrostatic pressure relaxations with the root pressure probe are commonly used for measuring the hydraulic conductivity (Lp(r)) of roots. We compared the Lp(r) of roots from species with different root hydraulic properties (Lupinus angustifolius L. 'Merrit', Lupinus luteus L. 'Wodjil', Triticum aestivum L. 'Kulin' and Zea mays L. 'Pacific DK 477') using pressure relaxations, a pressure clamp and osmotic gradients to induce water flow across the root. Only the pressure clamp measures water flow under steady-state conditions. Lp(r) determined by pressure relaxations was two- to threefold greater than Lp(r) from pressure clamps and was independent of the direction of water flow. Lp(r) (pressure clamp) was two- to fourfold higher than for Lp(r) (osmotic) for all species except Triticum aestivum where Lp(r) (pressure clamp) and Lp(r) (osmotic) were not significantly different. A novel technique was developed to measure the propagation of pressure through roots to investigate the cause of the differences in Lp(r). Root segments were connected between two pressure probes so that when root pressure (P(r)) was manipulated by one probe, the other probe recorded changes in P(r). Pressure relaxations did not induce the expected kinetics in pressure in the probe at the other end of the root when axial hydraulic conductance, and probe and root capacitances were accounted for. An electric circuit model of the root was constructed that included an additional capacitance in the root loaded by a series of resistances. This accounted for the double exponential kinetics for intact roots in pressure relaxation experiments as well as the reduced response observed with the double probe experiments. Although there were potential errors with all the techniques, we considered that the measurement of Lp(r) using the pressure clamp was the most unambiguous for small pressure changes, and provided that sufficient time was allowed for pressure propagation through the root. The differences in Lp(r) from different methods of measurement have implications for the models describing water transport through roots and the potential role of aquaporins.

The water transport system in plants provides a great and largely unexplored potential for improving the performance of crop plants. Water supply is well known as a critical limitation to crop productivity, but less well known is the fact that a substantial limitation to water supply occurs within the plant, due to a finite capacity of roots, stems and leaves to transport water. Understanding the limitations to maximum water transport in crop plants and what regulates the dynamic permeability of plant tissues provides an opportunity to enhance carbon accumulation and growth by ensuring sufficient water supply. In this chapter we examine the principles of plant water transport, with particular focus on how the anatomy of the xylem water-delivery system in the roots and leaves affects the efficiency with which water is supplied for transpiration under ample and water-deficit soil conditions, and present a vascular-centric view of the potential for improving crop yield.

Dieback of riparian species on floodplains has been attributed to increased soil salinisation due to raised groundwater levels, resulting from irrigation and river regulation. This is exacerbated by a reduction in flooding frequency and duration of inundation. For the Chowilla floodplain on the River Murray raised water tables have increased the amount of salts mobilised in the soil profile, causing the trees to experience salt induced water stress. For the trees to survive in the long term, salts need to be leached from the root zone. This study investigated whether floodwater infiltrates through channels created by E. largiflorens (black box) roots, flushing salts away from roots, thereby allowing the trees to increase their water uptake. Trees at different sites on the floodplain were artificially flooded, by pumping 1.5 kL of creek water into impoundments constructed around the trees. Gas exchange parameters, and pre-dawn and midday water potential were measured the day before, the day after and one week after the artificial flood and compared against trees that were not flooded. Pre-dawn and midday water potentials were also measured one month after the flood. After flooding, the trees experienced less water stress, indicated by an increase in water potential of less than 0.2 MPa, in comparison to non-flooded control trees. However, this response was not evident one month after flooding. The response to flooding did not result in increased rates of transpiration, stomatal conductance or photosynthesis, even though flooding effectively doubled the trees yearly water supply. The infiltration of floodwater in the impoundments around E. largiflorens was also compared to that of impoundments on bare ground. Floodwater infiltrated 2 – 17 times faster around trees than on adjacent bare ground, for parts of the floodplain not grazed by livestock. Tracer dye experiments indicated that bulk flow of water through pores down the profile was the reason for the enhanced infiltration. Flooding leached salts in direct vicinity of tree roots, but only leached small amounts of salts from the bulk soil.

There is strong evidence that aquaporins are central components in plant water relations. Plant species possess more aquaporin genes than species from other kingdoms. According to sequence similarities, four major groups have been identified, which can be further divided into subgroups that may correspond to localization and transport selectivity. They may be involved in compatible solute distribution, gas-transfer (CO2, NH3) as well as in micronutrient uptake (boric acid). Recent advances in determining the structure of some aquaporins gives further details on the mechanism of selectivity. Gating behaviour of aquaporins is poorly understood but evidence is mounting that phosphorylation, pH, pCa and osmotic gradients can affect water channel activity. Aquaporins are enriched in zones of fast cell division and expansion, or in areas where water flow or solute flux density would be expected to be high. This includes biotrophic interfaces between plants and parasites, between plants and symbiotic bacteria or fungi, and between germinating pollen and stigma. On a cellular level aquaporin clusters have been identified in some membranes. There is also a possibility that aquaporins in the endoplasmic reticulum may function in symplasmic transport if water can flow from cell to cell via the desmotubules in plasmodesmata. Functional characterization of aquaporins in the native membrane has raised doubt about the conclusiveness of expression patterns alone and need to be conducted in parallel. The challenge will be to elucidate gating on a molecular level and cellular level and to tie those findings into plant water relations on a macroscopic scale where various flow pathways need to be considered.

This chapter discusses the impacts of climate change on agriculture and the role of adaptation. Specifically, the chapter explores the following impacts of climate change on agriculture: increased CO2 concentrations, water availability, and pests and diseases. The chapter also includes reports of case studies on climate change impacts on wheat, rice and millet.

Terminal drought is a common abiotic stress affecting wheat yield in Mediterranean-type environments. As terminal drought develops, top layers of the soil profile dry, exposing the upper part of the root system to soil water deficit while deeper roots can still access soil water. Since open stomata rapidly exhausts available soil water, reducing stomatal conductance to prolong availability of soil water during grain filling may improve wheat yields in water-limited environments. It was hypothesised that genotypes with more root biomass in the drying upper layer of the soil profile accumulate more abscisic acid in the leaf and initiate stomatal closure to regulate water use under terminal drought. The wheat cultivar Drysdale and the breeding line IGW-3262 were grown in pots horizontally split into two segments by a wax-coated layer that hydraulically isolated the top and bottom segments, but allowed roots to grow into the bottom segment. Terminal drought was induced from anthesis by withholding water from (i) the top segment only (DW) and (ii) the top and bottom segments (DD) while both segments in well-watered pots (WW) were maintained at 90% pot soil water capacity. Drysdale, initiated stomatal closure earlier than IGW-3262, possibly due to higher signal strength generated in its relatively larger proportion of roots in the drying top segment. The relationship between leaf ABA and stomatal conductance was strong in Drysdale but weak in IGW-3262. Analysis of ABA metabolites suggests possible differences in ABA metabolism between these two genotypes. A higher capability of deeper roots to extract available water is also important in reducing the gap between actual and potential yield.

Wheat (Triticum aestivum L.) production may be affected by the future climate, but the impact of the combined increases in atmospheric CO2 concentration, temperature and incidence of drought that are predicted has not been evaluated. The combined effect of elevated CO2, high temperature and terminal drought on biomass accumulation and grain yield was evaluated in vigorous (38-19) and nonvigorous (Janz) wheat genotypes grown under elevated CO2 (700µLL-1) combined with temperatures 2°C, 4°C and 6°C above the current ambient temperature. Terminal drought was induced in all combinations at anthesis in a split-plot design to test whether the effect of elevated CO2 combined with high temperature ameliorates the negative effects of terminal drought on biomass accumulation and grain yield. Biomass and grain yield were enhanced under elevated CO2 with 2°C above the ambient temperature, regardless of the watering regimen. The combinations of elevated CO2 plus 4°C or 6°C above the ambient temperature did not enhance biomass and grain yield, but tended to decrease them. The reductions in biomass and grain yield (45-50%) caused by terminal drought were less severe (21-28%) under elevated CO2 with 2°C above the ambient temperature. The amelioration resulted from a 63% increase in the rate of leaf net photosynthesis in 38-19 and a 39% increase in tillering and leaf area in Janz. The contrasting responses and phenological development of these two genotypes to the combination of elevated CO2, temperature and terminal drought, and the possible influences on their source-sink relationships are discussed.

The ‘Old World’ lupin species are unique among grain legume crops in their strong specific adaption to acid sandy soils, and in their extremely recent domestication history. Our understanding of lupin responses to drought is limited; based on studies with elite cultivars, representing only a small fraction of the genetic diversity resident in the species, and subjected to strong selection for early phenology throughout their domesticated history, facilitating drought escape. Lupins appear to have a mix of competitive and conservative water use strategies, with profligate water use and high rates of photosynthesis when water is freely available; coupled with high sensitivity to water deficits, whereby stomatal conductance is reduced, phytohormone concentrations and metabolism modified, well before changes in leaf water potential occur. With high root-shoot ratios and high hydraulic conductance, lupin roots are highly efficient at taking up and transporting water. However, the predominantly apoplastic flow in lupins cannot be regulated as flexibly as the symplastic water transport that is so important in cereals. Low rates of remobilization of pre-anthesis stored C in lupin forces the crop to rely almost completely on current photosynthesis, which exacerbates the effects of terminal drought, given that the species respond by reducing leaf area through abscission. Because of the past narrow focus on domesticated material it is not currently possible to put these observations in an ecophysiological context, to answer which of these attributes are characteristic of lupins as a species, and which can be expected to vary in response to environmental selection pressure. To advance our understanding of the species we advocate the study of wild germplasm specifically-adapted to habitats that impose contrasting drought stress, to address both the narrowness and short evolutionary history of the domesticated material, by highlighting responses to millennia of natural selection. By identifying the pros and cons of adaptive traits in an ecophysiological context our capacity to improve elite material will be considerably advanced.

In south-west Australia, winter grown crops such as wheat and lupin often experience transient waterlogging during periods of high rainfall. Wheat is believed to be more tolerant to waterlogging than lupins, but until now no direct comparisons have been made. The effects of waterlogging on root growth and anatomy were compared in wheat (Triticum aestivum L.), narrow-leafed lupin (Lupinus angustifolius L.) and yellow lupin (Lupinus luteus L.) using 1 m deep root observation chambers. Seven days of waterlogging stopped root growth in all species, except some nodal root development in wheat. Roots of both lupin species died back progressively from the tips while waterlogged. After draining the chambers, wheat root growth resumed in the apical region at a faster rate than well-drained plants, so that total root length was similar in waterlogged and well-drained plants at the end of the experiment. Root growth in yellow lupin resumed in the basal region, but was insufficient to compensate for root death during waterlogging. Narrow-leafed lupin roots did not recover; they continued to deteriorate. The survival and recovery of roots in response to waterlogging was related to anatomical features that influence internal oxygen deficiency and root hydraulic properties.

The hydraulic properties of plant roots depend on the morphology and anatomy of the root system, the length of the absorbing region and the influence of aquaporins (AQPs). These features change during development and in response to environmental stimuli, and alter the hydraulic conductivity of the root system ( Lp r ). AQPs are proteins that form water selective channels to facilitate water flow across membranes. A large proportion of AQP isoforms are predominantly expressed in roots and their localization indicates a putative role in the transport of water across the root. AQP activity can finely regulate the rate of water flow across the root by changes in abundance and opening/closing the water channels. Since water will flow by the pathway of least resistance, AQPs will only influence radial water flow if the hydraulic conductivity of the apoplast is relatively less than that of the cell‐to‐cell pathway. There is growing evidence that AQPs influence water flow through the roots of some, but not all, species. Waterlogging is a significant environmental constraint to crop growth, but its influence on Lp r is poorly understood. Depending on the tolerance of the species, waterlogging through oxygen deficiency reduces root growth and tends to reduce Lp r . Oxygen deficiency can directly or indirectly close AQPs or alter their abundance. Changes in AQP activity may be the key component which ultimately influences water transport through waterlogged roots.

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Wheat (Triticum aestivum L.) production may be affected by the future climate, but the impact of the combined increases in atmospheric CO2 concentration, temperature and incidence of drought that are predicted has not been evaluated. The combined effect of elevated CO2, high temperature and terminal drought on biomass accumulation and grain yield was evaluated in vigorous (38–19) and nonvigorous (Janz) wheat genotypes grown under elevated CO2 (700 µL L–1) combined with temperatures 2°C, 4°C and 6°C above the current ambient temperature. Terminal drought was induced in all combinations at anthesis in a split-plot design to test whether the effect of elevated CO2 combined with high temperature ameliorates the negative effects of terminal drought on biomass accumulation and grain yield. Biomass and grain yield were enhanced under elevated CO2 with 2°C above the ambient temperature, regardless of the watering regimen. The combinations of elevated CO2 plus 4°C or 6°C above the ambient temperature did not enhance biomass and grain yield, but tended to decrease them. The reductions in biomass and grain yield (45–50%) caused by terminal drought were less severe (21–28%) under elevated CO2 with 2°C above the ambient temperature. The amelioration resulted from a 63% increase in the rate of leaf net photosynthesis in 38–19 and a 39% increase in tillering and leaf area in Janz. The contrasting responses and phenological development of these two genotypes to the combination of elevated CO2, temperature and terminal drought, and the possible influences on their source–sink relationships are discussed.

Little is known about water flow across intact root cells and roots in response to hypoxia. Responses may be rapid if regulated by aquaporin activity, but only if water crosses membranes. We measured the transport properties of roots and cortical cells of three important crop species in response to hypoxia (0.05 mol O2 m–3): wheat (Triticum aestivum L.), narrow-leafed lupin (Lupinus angustifolius L.) and yellow lupin (Lupinus luteus L.). Hypoxia influenced solute transport within minutes of exposure as indicated by increases in root pressure (Pr) and decreases in turgor pressure (Pc), but these effects were only significant in lupins. Re-aeration returned Pr to original levels in yellow lupin, but in narrow-leafed lupin, Pr declined to zero or lower values without recovery even when re-aerated. Hypoxia inhibited hydraulic conductivity of root cortical cells (Lpc) in all three species, but only inhibited hydraulic conductivity of roots (Lpr) in wheat, indicating different pathways for radial water flow across lupin and wheat roots. The inhibition of Lpr of wheat depended on the length of the root, and inhibition of Lpc in the endodermis could account for the changes in Lpr. During re-aeration, aquaporin activity increased in wheat roots causing an overshoot in Lpr. The results of this study demonstrate that the roots of these species not only vary in hydraulic properties but also vary in their sensitivity to the same external O2 concentration.