Silicon fertilisation affects morphological and immune defences of an insect pest and enhances plant compensatory growth

Herbivorous insects have evolved various anti-predator defences, including morphological, behavioural, and immune defences, which can make biocontrol of herbivorous pests challenging. Silicon (Si) accumulation in plants is a potent physical defence against mandibulate insects. However, it remains uncertain how Si affects the anti-predator defences of insect herbivores and plant defences following herbivory. We grew the model grass, Brachypodium distachyon, hydroponically with (+Si) or without (-Si) Si and investigated the plant-mediated effects of Si on the anti-predator defences of the cotton bollworm, Helicoverpa armigera, integrating morphological (i.e. integument resistance and thickness), behavioural, and immune defences. We also examined the effects of Si on plant compensatory growth and leaf trichome production. Larval growth, leaf consumption, and integument resistance were lower when feeding on +Si plants compared to when feeding on -Si plants. Larval integument thickness, defensive behaviours, haemocyte density, and lysozyme-like activity in the haemolymph were unaffected by Si. Larvae fed on +Si plants had higher haemolymph phenoloxidase (PO) and total-PO activities than larvae fed on -Si plants, although this did not enhance the melanisation response of larvae. Furthermore, Si supplies increased plant compensation for herbivory and constitutive trichome production, whereas herbivory induced trichome production only on -Si plants. We provide the first evidence for plant-mediated effects of Si on anti-predator defences of an insect herbivore. We suggest that the lower integument resistance of larvae when feeding on Si-supplemented plants could contribute to their vulnerability to natural enemies and that high PO activity may impose fitness costs (e.g. delayed development).