Abstract
Insect herbivores employ various defences, including morphological, behavioural, and immune responses against their natural enemies (e.g., predators, parasitoids) which can make biocontrol of herbivorous pests challenging. Silicon (Si) accumulation in plants is a potent physical defence against herbivores. However, it remains uncertain how Si affects pest defences to their enemies and plant defences following herbivore attack. We grew the model grass, Brachypodium distachyon, hydroponically with (+Si) or without (–Si) Si and investigated the impacts of Si on morphological (integument resistance and thickness), behavioural (flee, headrear, thrash, and regurgitation), and immune defences of the cotton bollworm, Helicoverpa armigera. We further 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, hemocyte density and lysozyme-like activity in the hemolymph were unaffected by Si. Larvae fed on +Si plants had higher hemolymph phenoloxidase (PO) and total-PO activities than larvae fed on –Si plants, although this did not enhance larval melanisation response. Furthermore, Si supply increased plant compensatory growth and constitutive trichome production whereas herbivory induced trichome production only on –Si plants. We provide the first evidence that Si fertilisation affects insect defences in addition to reducing their growth and feeding. Lower integument resistance might enhance larval vulnerability to parasitoids and pathogens and higher PO activities could impose fitness costs (e.g., delayed development), potentially increasing overall insect susceptibility to enemies.
Memory of the vernalized state in plants including the model grass Brachypodium distachyon