Climatic Drivers of Silicon Accumulation in a Model Grass Operate in Low- but Not High-Silicon Soils.

Grasses are hyper-accumulators of silicon (Si), which is known to alleviate diverse environmental stresses, prompting speculation that Si accumulation evolved in response to unfavourable climatic conditions, including seasonally arid environments. We conducted a common garden experiment using 57 accessions of the model grass Brachypodium distachyon, sourced from different Mediterranean locations, to test relationships between Si accumulation and 19 bioclimatic variables. Plants were grown in soil with either low or high (Si supplemented) levels of bioavailable Si. Si accumulation was negatively correlated with temperature variables (annual mean diurnal temperature range, temperature seasonality, annual temperature range) and precipitation seasonality. Si accumulation was positively correlated with precipitation variables (annual precipitation, precipitation of the driest month and quarter, and precipitation of the warmest quarter). These relationships, however, were only observed in low-Si soils and not in Si-supplemented soils. Our hypothesis that accessions of B. distachyon from seasonally arid conditions have higher Si accumulation was not supported. On the contrary, higher temperatures and lower precipitation regimes were associated with lower Si accumulation. These relationships were decoupled in high-Si soils. These exploratory results suggest that geographical origin and prevailing climatic conditions may play a role in predicting patterns of Si accumulation in grasses.

Nitrogen availability and allocation in sorghum and its wild relatives: Divergent roles for cyanogenic glucosides.

Crop plants are assumed to have become more susceptible to pests as a result of selection for high growth rates during the process of domestication, consistent with resource allocation theories. We compared the investment by domesticated sorghum into cyanogenic glucosides, nitrogen-based specialised metabolites that break down to release hydrogen cyanide, with five wild relatives native to Australia. Plants were grown in pots in a greenhouse and supplied with low and high concentrations of nitrogen and monitored for 9 weeks. The concentrations of nitrate, total phenolics and silicon were also measured. Domesticated Sorghum bicolor had the highest leaf and root cyanogenic glucoside concentrations, and among the lowest nitrate and silicon concentrations under both treatments. Despite partitioning a much higher proportion of its stored nitrogen to cyanogenic glucosides than the wild species, S. bicolor's nitrogen productivity levels were among the highest. Most of the wild sorghums had higher concentrations of silicon and phenolics, which may provide an alternative defence system. Cyanogenic glucosides appear to be integral to S. bicolor's physiology, having roles in both growth and defence. Sorghum macrospermum displayed consistently low cyanogenic glucoside concentrations, high growth rates and high nitrogen productivity and represents a particularly attractive genetic resource for sorghum improvement.

Silicon deposition on guard cells increases stomatal sensitivity as mediated by K+ efflux and consequently reduces stomatal conductance.

Silicon (Si) has been widely reported to improve plant resistance to water stress via various mechanisms including cuticular Si deposition to reduce leaf transpiration. However, there is limited understanding of the effects of Si on stomatal physiology, including the underlying mechanisms and implications for resistance to water stress. We grew tall fescue (Festuca arundinacea Schreb. cv. Fortuna) hydroponically, with or without Si, and treated half of the plants with 20% polyethylene glycol to impose physiological drought (osmotic stress). Scanning electron microscopy in conjunction with X-ray mapping found that Si was deposited on stomatal guard cells and as a sub-cuticular layer in Si-treated plants. Plants grown in Si had a 28% reduction in stomatal conductance and a 23% reduction in cuticular conductance. When abscisic acid was applied exogenously to epidermal leaf peels to promote stomatal closure, Si plants had 19% lower stomatal aperture compared to control plants (i.e. increased stomatal sensitivity) and an increased efflux of guard cell K+ ions. However, the changes in stomatal physiology with Si were not substantial enough to improve water stress resistance, as shown by a lack of significant effect of Si on water potential, growth, photosynthesis and water-use efficiency. Our findings suggest a novel underlying mechanism for reduced stomatal conductance with Si application; specifically, that Si deposition on stomatal guard cells promotes greater stomatal sensitivity as mediated by guard cell K+ efflux.

The Role of Silicon in Antiherbivore Phytohormonal Signalling.

The role of plant silicon (Si) in the alleviation of abiotic and biotic stress is now widely recognised and researched. Amongst the biotic stresses, Si is known to increase resistance to herbivores through biomechanical and chemical mechanisms, although the latter are indirect and remain poorly characterised. Chemical defences are principally regulated by several antiherbivore phytohormones. The jasmonic acid (JA) signalling pathway is particularly important and has been linked to Si supplementation, albeit with some contradictory findings. In this Perspectives article, we summarise existing knowledge of how Si affects JA in the context of herbivory and present a conceptual model for the interactions between Si and JA signalling in wounded plants. Further, we use novel information from the model grass Brachypodium distachyon to underpin aspects of this model. We show that Si reduces JA concentrations in plants subjected to chemical induction (methyl jasmonate) and herbivory (Helicoverpa armigera) by 34% and 32%, respectively. Moreover, +Si plants had 13% more leaf macrohairs than -Si plants. From this study and previous work, our model proposes that Si acts as a physical stimulus in the plant, which causes a small, transient increase in JA. When +Si plants are subsequently attacked by herbivores, they potentially show a faster induction of JA due to this priming. +Si plants that have already invested in biomechanical defences (e.g. macrohairs), however, have less utility for JA-induced defences and show lower levels of JA induction overall.

Barley yellow dwarf virus infection and elevated CO2 alter the antioxidants ascorbate and glutathione in wheat.

Plant antioxidants ascorbate and glutathione play an important role in regulating potentially harmful reactive oxygen species produced in response to virus infection. Barley yellow dwarf virus is a widespread viral pathogen that systemically infects cereal crops including wheat, barley and oats. In addition, rising atmospheric CO2 will alter plant growth and metabolism, including many potential but not well understood effects on plant-virus interactions. In order to better understand the wheat-BYDV interaction and any potential changes under elevated CO2, the total concentration and oxidised fraction of ascorbate and glutathione was measured in leaves of a susceptible wheat cultivar (Triticum aestivum L. 'Yitpi') infected with Barley yellow dwarf virus-PAV (Padi Avenae virus) and grown under elevated CO2 in controlled environment chambers. Virus infection decreased total leaf ascorbate and glutathione concentrations and increased the fraction of oxidised ascorbate (dehydroascorbate). Elevated CO2 decreased the fraction of oxidised ascorbate. In this work, we demonstrate that systemic infection by a phloem-restricted virus weakens the antioxidant pools of ascorbate and glutathione. In addition, elevated CO2 may decrease oxidative stress, for example, from virus infection, but there was no direct evidence for an interactive effect between treatments.

Virus infection mediates the effects of elevated CO2 on plants and vectors.

Atmospheric carbon dioxide (CO2) concentration has increased significantly and is projected to double by 2100. To increase current food production levels, understanding how pests and diseases respond to future climate driven by increasing CO2 is imperative. We investigated the effects of elevated CO2 (eCO2) on the interactions among wheat (cv. Yitpi), Barley yellow dwarf virus and an important pest and virus vector, the bird cherry-oat aphid (Rhopalosiphum padi), by examining aphid life history, feeding behavior and plant physiology and biochemistry. Our results showed for the first time that virus infection can mediate effects of eCO2 on plants and pathogen vectors. Changes in plant N concentration influenced aphid life history and behavior, and N concentration was affected by virus infection under eCO2. We observed a reduction in aphid population size and increased feeding damage on noninfected plants under eCO2 but no changes to population and feeding on virus-infected plants irrespective of CO2 treatment. We expect potentially lower future aphid populations on noninfected plants but no change or increased aphid populations on virus-infected plants therefore subsequent virus spread. Our findings underscore the complexity of interactions between plants, insects and viruses under future climate with implications for plant disease epidemiology and crop production.

Drought adversely affects tuber development and nutritional quality of the staple crop cassava (Manihot esculenta Crantz).

Cassava (Manihot esculenta Crantz) is the staple food source for over 850million people worldwide. Cassava contains cyanogenic glucosides and can be toxic to humans, causing paralysing diseases such as konzo, and even death if not properly processed. Konzo epidemics are often associated with times of drought. This may be due to a greater reliance on cassava as it is drought tolerant, but it may also be due to an increase in cyanogenic glucosides. Episodic droughts are forecast to become more common in many cassava-growing regions. We therefore sought to quantify the effect of water-stress on both yield and cyanogenic glucoside concentration (CNc) in the developing tubers of cassava. Five-month-old plants were grown in a glasshouse and either well watered or droughted for 28 days. A subset of droughted plants was re-watered half way through the experiment. Droughted plants had 45% fewer leaves and lower tuber yield, by 83%, compared with well-watered plants. CNc was 2.9-fold higher in the young leaves of droughted plants, whereas CNc in tubers from droughted plants was 4-fold greater than in tubers from well-watered plants. Re-watered plants had a similar biomass to control plants, and lower CNc than droughted plants. These findings highlight the important link between food quality and episodic drought.