Helen Bramley

Heat Stress in Wheat during Reproductive and Grain-Filling Phases

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 physiological and biotechnological tools with conventional breeding techniques will help to develop wheat varieties with better grain yield under heat stress during reproductive and grain-filling phases. This review discusses the impact of heat stress during reproductive and grain-filling stages of wheat on grain yield and suggests strategies to improve heat stress tolerance in wheat.

Roles of Morphology, Anatomy, and Aquaporins in Determining Contrasting Hydraulic Behavior of Roots

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 (Lax) and the distribution of water uptake along the root, with a more localized role for aquaporins (AQPs). Lupin roots had greater Lax than wheat roots, due to greater xylem development. Lax and root hydraulic conductance (Lr) were related to each other, such that both variables increased with distance from the root tip in lupin roots. Lax and Lr were constant with distance from the tip in wheat roots. Despite these contrasting behaviors, the hydraulic conductivity of root cells (Lpc) was similar for all species and increased from the root surface toward the endodermis. Lpc was largely controlled by AQPs, as demonstrated by dramatic reductions in Lpc 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 Lr. This study demonstrates the importance of examining root morphology and anatomy in assessing the role of AQPs in root hydraulics.

Neglecting legumes has compromised human health and sustainable food production

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-used. 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.

Water Flow in the Roots of Crop Species: The Influence of Root Structure, Aquaporin Activity, and Waterlogging

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.

The contrasting influence of short-term hypoxia on the hydraulic properties of cells and roots of wheat and lupin

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.

Floodwater infiltration through root channels on a sodic clay floodplain and the influence on a local tree species Eucalyptus largiflorens

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.

Non-invasive pressure probes magnetically clamped to leaves to monitor the water status of wheat

Background and aimsBeing 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.MethodsThe probes consist of miniature pressure sensors that clamp to the leaves via magnets and detect relative changes in hydration status. Probes were clamped to leaves of six individual plants of the cultivar Wyalkatchem at the stem elongation stage and compared against traditional plant water relations measurements.ResultsOutput from the probes, called patch-pressure (Pp), correlated well with leaf water potential and transpiration of individual plants. Variation between plants in the original clamp pressure exerted by the magnets and leaf individual properties led to variations in the amplitude of the diurnal Pp profiles, but not in the kinetics of the curves where Pp responded simultaneously in all plants to changes in the ambient environment (light and temperature).ConclusionsDrying and rewatering cycles and analysis of the curve kinetics identified several methods that can be used to test comparisons of water status monitoring of wheat genotypes under water deficit.

Root growth of lupins is more sensitive to waterlogging than wheat

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 1m 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.

Root water transport under waterlogged conditions and the roles of aquaporins

Water flow through plants roots can be affected when the soil is waterlogged and oxygen deficient. For species not adapted to these conditions, water flow usually decreases within minutes to days, depending on the oxygen concentration in the root and rhizosphere. During this time, the decrease in water flow is attributed to decreased root hydraulic conductance, through an inhibition of plasma-membrane aquaporins. There is increasing evidence that aquaporins may also be involved in the transport of gases, end products of anaerobic respiration, and signalling molecules; all of which are relevant to oxygen-deficient conditions. Eventually, primary roots die if continually starved of oxygen, but may be replaced with adventitious roots that can maintain the supply of water to the shoot. Here, we review the effects of waterlogging and oxygen deficiency on root hydraulic conductance and aquaporin activity.

Response of wheat restricted-tillering and vigorous growth traits to variables of climate change.

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.