Leaf Silicification Provides Herbivore Defence Regardless Of The Extensive Impacts Of Water Stress

FUNCTIONAL ECOLOGY(2021)

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Abstract
Altered precipitation patterns due to climate change are likely to impose water-deficit stress in plants resulting in changes to specific leaf mass, leaf water content and chemical defences that may impact herbivorous arthropods. Grasses, in particular, accumulate large concentrations of silicon (Si) which provides physical defence against herbivores. Although Si uptake by plants may be affected by water availability, very few studies have investigated the combined effect of water-deficit stress and Si on insect herbivore performance.We grew tall fescue Festuca arundinacea Schreb. hydroponically, with and without Si, and half of the plants were treated with 20% polyethylene glycol (PEG) to impose osmotic stress. In all, 11 leaf traits (physiological, chemical and structural) were measured, silicified phytoliths on the leaf surface were visualised using scanning electron microscopy (SEM) in conjunction with X-ray mapping, and plants were exposed to a chewing insect herbivore [Helicoverpa armigera Hubner (Lepidoptera: Noctuidae)].Although osmotic stress was associated with changes to leaf physiological and chemical traits, including increased specific leaf mass, decreased leaf relative water content and increased leaf nitrogen (N), there was no significant effect on H. armigera relative growth rate (RGR). However, Si reduced RGR of H. armigera by 80%-98% while generating few changes to physiological and chemical leaf traits. Instead, the decline in RGR with Si was associated with changes to leaf structural traits, in particular, a greater density of silicified phytoliths on the leaf surface.Comparison of effect sizes indicated that leaf traits were primarily affected by osmotic stress but not Si, and that herbivore RGR was strongly negatively affected by Si but not osmotic stress. There was no interactive effect between the osmotic stress and Si treatments on H. armigera RGR or plant traits except for leaf nitrogen and phenolic concentrations. This study provides further support that Si may prove to be beneficial to plants against chewing insect pests and remains robust regardless of water-deficit stress conditions.A free Plain Language Summary can be found within the Supporting Information of this article.
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Key words
drought, insect herbivory, nutritional quality, osmotic stress, physical defence, plant traits, silicon, water-deficit stress
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