William J. Davies

Drought, ozone, ABA and ethylene: new insights from cell to plant to community.

Recent reports show ethylene-dependent reductions in stomatal sensitivity to abscisic acid (ABA) under ozone stress. These changes reduce stomatal control of plant water loss in drying soil. Here we review evidence that ABA and ethylene, and interactions between these two stress-induced hormones, control many of the responses of intact plants to drought and ozone stress, with emphasis on effects on stomata and shoot growth. We draw attention to convergent signalling and response pathways induced by ozone and drought that can increase production of hydrogen peroxide (H(2)O(2)) and nitric oxide (NO). Stomatal responses to a wider range of stresses and developmental cues may also be controlled via the same sets of signalling pathways. Other hormones, or effectors such as xylem/apoplastic pH or changes in plant water status, also play a role in signalling within and between organs. We discuss the implications, for crops, natural ecosystems and water catchment processes, of ethylenes antagonism of the stomatal response to ABA, against a back-drop of predictions for reduced precipitation and increasing ozone pollution, as part of global climate change and increasing urbanization and industrial development.

Assessing shifts in microbial community structure across a range of grasslands of differing management intensity using CLPP, PLFA and community DNA techniques.

This study aimed to characterise soil microbial community structure and function in temperate upland grassland ecosystems. We compared the use of community level physiological profiles (CLPP), phospholipid fatty acid (PLFA) profiles and community DNA (%G+C base distribution) approaches to quantify soil microbial community structure and potential activity across a gradient of three upland grassland types at 10 geographically distinct sites within the UK. Soil microbial biomass (Cmic) was highest in unimproved (U4a) and lowest in improved (MG6) grasslands. In contrast, potential soil microbial activity (carbon utilisation) was greatest in the improved and lowest in the semi-improved (U4b) grasslands. PLFA and culturing revealed that the soil microbial community shifted from one favouring fungi to one favouring bacteria as grassland improvement increased. Canonical variate analysis (CVA) of the CLPP and PLFA data differentiated microbial communities from the grassland types and sites and the separation between grasslands was greater using PLFA than CLPP. Discrimination between grasslands was mainly due to the presence of higher concentrations of fatty acids typical for Gram −ve bacteria in improved grasslands and actinomycete and fungal fatty acids in the semi and unimproved grasslands. CVA of the %G+C data gave less discrimination of the microbial communities than the other two methods. Correlation analysis of the CVA data for each microbial analysis showed a small, but significant, level of matching between the CLPP and PLFA data suggesting these two analyses may be reporting on similar members of the microbial community. Correlation between microbial community structure and soil physio-chemical properties indicated that PLFA were highly correlated with calcium, phosphorus, sodium, nitrogen and organic matter content and pH. CLPP were highly correlated with sodium and organic matter content and pH, while %G+C content correlated with pH. Correlation between microbial community structure and plant community structure indicated that fatty acids typical for Gram −ve bacteria were highly correlated with the presence of Lolium perenne and Trifolium repens and all microbial PLFA with the presence of Vaccinium myrtillus. Correlation of plant species with CLPP indicated that the presence of a number of rushes, shrubs, herbs and grasses influenced the metabolic profiles of the microbial communities from these grasslands. The presence of herbs were found to be highly correlated with certain %G+C classes within the community DNA.

Improving water use in crop production.

Globally, agriculture accounts for 80–90% of all freshwater used by humans, and most of that is in crop production. In many areas, this water use is unsustainable; water supplies are also under pressure from other users and are being affected by climate change. Much effort is being made to reduce water use by crops and produce ‘more crop per drop’. This paper examines water use by crops, taking particularly a physiological viewpoint, examining the underlying relationships between carbon uptake, growth and water loss. Key examples of recent progress in both assessing and improving crop water productivity are described. It is clear that improvements in both agronomic and physiological understanding have led to recent increases in water productivity in some crops. We believe that there is substantial potential for further improvements owing to the progress in understanding the physiological responses of plants to water supply, and there is considerable promise within the latest molecular genetic approaches, if linked to the appropriate environmental physiology. We conclude that the interactions between plant and environment require a team approach looking across the disciplines from genes to plants to crops in their particular environments to deliver improved water productivity and contribute to sustainability.

Xylem Sap pH Increase: A Drought Signal Received at the Apoplastic Face of the Guard Cell That Involves the Suppression of Saturable Abscisic Acid Uptake by the Epidermal Symplast

Drought increased the pH of Commelina communis xylem sap from 6.1 to 6.7. Conductances of transpiring leaves were 50% lower in pH 7.0 than in pH 6.0 buffers, but bulk leaf abscisic acid (ABA) concentration and shoot water status were unaffected by pH. Stomatal apertures of isolated abaxial epidermis incubated on simple buffers increased with external pH, so in vivo this must be overridden by alternative pH effects. Reductions in leaf transpiration rate at pH 7.0 were dependent on the presence of 10–8 mol dm-3 ABA in the xylem stream. We inferred that at pH 7.0 leaf apoplastic ABA concentrations increased: pH did not affect distributions of ABA among leaf tissues, but isolated epidermis and mesophyll tissue took up more 3H-ABA from pH 6.0 than from pH 7.0 buffers. The apoplastic ABA increase at pH 7.0 may result from reduced symplastic sequestration. A portion of 3H-ABA uptake by the epidermis was saturable at pH 6.0 but not at pH 7.0. An ABA uptake carrier may contribute to ABA sequestration by the leaf symplast of well-watered plants, and its inactivity at pH 7.0 may favor apoplastic ABA accumulation in draughted plants. Effects of external pH on stomatal apertures in the isolated epidermis indicate that published data supporting a role for internal guard cell ABA receptors should be reassessed.

Raising yield potential of wheat. III. Optimizing partitioning to grain while maintaining lodging resistance

A substantial increase in grain yield potential is required, along with better use of water and fertilizer, to ensure food security and environmental protection in future decades. For improvements in photosynthetic capacity to result in additional wheat yield, extra assimilates must be partitioned to developing spikes and grains and/or potential grain weight increased to accommodate the extra assimilates. At the same time, improvement in dry matter partitioning to spikes should ensure that it does not increase stem or root lodging. It is therefore crucial that improvements in structural and reproductive aspects of growth accompany increases in photosynthesis to enhance the net agronomic benefits of genetic modifications. In this article, six complementary approaches are proposed, namely: (i) optimizing developmental pattern to maximize spike fertility and grain number, (ii) optimizing spike growth to maximize grain number and dry matter harvest index, (iii) improving spike fertility through desensitizing floret abortion to environmental cues, (iv) improving potential grain size and grain filling, and (v) improving lodging resistance. Since many of the traits tackled in these approaches interact strongly, an integrative modelling approach is also proposed, to (vi) identify any trade-offs between key traits, hence to define target ideotypes in quantitative terms. The potential for genetic dissection of key traits via quantitative trait loci analysis is discussed for the efficient deployment of existing variation in breeding programmes. These proposals should maximize returns in food production from investments in increased crop biomass by increasing spike fertility, grain number per unit area and harvest index whilst optimizing the trade-offs with potential grain weight and lodging resistance.

Long-distance ABA Signaling and Its Relation to Other Signaling Pathways in the Detection of Soil Drying and the Mediation of the Plant’s Response to Drought

In this article we review evidence for a variety of long-distance signaling pathways involving hormones and nutrient ions moving in the xylem sap. We argue that ABA has a central role to play, at least in root-to-shoot drought stress signaling and the regulation of functioning, growth, and development of plants in drying soil. We also stress the importance of changes in the pH of the leaf cell apoplast as influenced both by edaphic and climatic variation, as a regulator of shoot growth and functioning, and we show how changes in xylem and apoplastic pH can affect the way in which ABA regulates stomatal behavior and growth. The sensitivity to drought of the pH/ABA sensing and signaling mechanism is emphasized. This allows regulation of plant growth, development and functioning, and particularly shoot water status, as distinct from stress lesions in growth and other processes as a reaction to perturbations such as soil drying.

Rhizosphere bacteria containing 1‐aminocyclopropane‐1‐carboxylate deaminase increase yield of plants grown in drying soil via both local and systemic hormone signalling

Decreased soil water availability can stimulate production of the plant hormone ethylene and inhibit plant growth. Strategies aimed at decreasing stress ethylene evolution might attenuate its negative effects. An environmentally benign (nonchemical) method of modifying crop ethylene relations - soil inoculation with a natural root-associated bacterium Variovorax paradoxus 5C-2 (containing the enzyme 1-aminocyclopropane-1-carboxylate (ACC) deaminase that degrades the ethylene precursor ACC), was assessed with pea (Pisum sativum) plants grown in drying soil. Inoculation with V. paradoxus 5C-2, but not with a transposome mutant with massively decreased ACC deaminase activity, improved growth, yield and water-use efficiency of droughted peas. Systemic effects of V. paradoxus 5C-2 included an amplified soil drying-induced increase of xylem abscisic acid (ABA) concentration, but an attenuated soil drying-induced increase of xylem ACC concentration. A local bacterial effect was increased nodulation by symbiotic nitrogen-fixing bacteria, which prevented a drought-induced decrease in nodulation and seed nitrogen content. Successfully deploying a single bacterial gene in the rhizosphere increased yield and nutritive value of plants grown in drying soil, via both local and systemic hormone signalling. Such bacteria may provide an easily realized, economic means of sustaining crop yields and using irrigation water more efficiently in dryland agriculture.

Improving crop productivity and resource use efficiency to ensure food security and environmental quality in China

In recent years, agricultural growth in China has accelerated remarkably, but most of this growth has been driven by increased yield per unit area rather than by expansion of the cultivated area. Looking towards 2030, to meet the demand for grain and to feed a growing population on the available arable land, it is suggested that annual crop production should be increased to around 580 Mt and that yield should increase by at least 2% annually. Crop production will become more difficult with climate change, resource scarcity (e.g. land, water, energy, and nutrients) and environmental degradation (e.g. declining soil quality, increased greenhouse gas emissions, and surface water eutrophication). To pursue the fastest and most practical route to improved yield, the near-term strategy is application and extension of existing agricultural technologies. This would lead to substantial improvement in crop and soil management practices, which are currently suboptimal. Two pivotal components are required if we are to follow new trajectories. First, the disciplines of soil management and agronomy need to be given increased emphasis in research and teaching, as part of a grand food security challenge. Second, continued genetic improvement in crop varieties will be vital. However, our view is that the biggest gains from improved technology will come most immediately from combinations of improved crops and improved agronomical practices. The objectives of this paper are to summarize the historical trend of crop production in China and to examine the main constraints to the further increase of crop productivity. The paper provides a perspective on the challenge faced by science and technology in agriculture which must be met both in terms of increased crop productivity but also in increased resource use efficiency and the protection of environmental quality.

The High Light Response in Arabidopsis Involves ABA Signaling between Vascular and Bundle Sheath Cells

Previously, it has been shown that Arabidopsis thaliana leaves exposed to high light accumulate hydrogen peroxide (H2O2) in bundle sheath cell (BSC) chloroplasts as part of a retrograde signaling network that induces ASCORBATE PEROXIDASE2 (APX2). Abscisic acid (ABA) signaling has been postulated to be involved in this network. To investigate the proposed role of ABA, a combination of physiological, pharmacological, bioinformatic, and molecular genetic approaches was used. ABA biosynthesis is initiated in vascular parenchyma and activates a signaling network in neighboring BSCs. This signaling network includes the Gα subunit of the heterotrimeric G protein complex, the OPEN STOMATA1 protein kinase, and extracellular H2O2, which together coordinate with a redox-retrograde signal from BSC chloroplasts to activate APX2 expression. High light–responsive genes expressed in other leaf tissues are subject to a coordination of chloroplast retrograde signaling and transcellular signaling activated by ABA synthesized in vascular cells. ABA is necessary for the successful adjustment of the leaf to repeated episodes of high light. This process involves maintenance of photochemical quenching, which is required for dissipation of excess excitation energy.

The Identification of Genes Involved in the Stomatal Response to Reduced Atmospheric Relative Humidity

Stomatal pores of higher plants close in response to decreases in atmospheric relative humidity (RH). This is believed to be a mechanism that prevents the plant from losing excess water when exposed to a dry atmosphere and as such is likely to have been of evolutionary significance during the colonization of terrestrial environments by the embryophytes. We have conducted a genetic screen, based on infrared thermal imaging, to identify Arabidopsis genes involved in the stomatal response to reduced RH. Here we report the characterization of two genes, identified during this screen, which are involved in the guard cell reduced RH signaling pathway. Both genes encode proteins known to be involved in guard cell ABA signaling. OST1 encodes a protein kinase involved in ABA-mediated stomatal closure while ABA2 encodes an enzyme involved in ABA biosynthesis. These results suggest, in contrast to previously published work, that ABA plays a role in the signal transduction pathway connecting decreases in RH to reductions in stomatal aperture. The identification of OST1 as a component required in stomatal RH and ABA signal transduction supports the proposition that guard cell signaling is organized as a network in which some intracellular signaling proteins are shared among different stimuli.