Plants are subjected to a multitude of stimuli during insect herbivory, resulting in a complex and cumulative defence response. Breaking down the components of herbivory into specific stimuli and identifying the mechanisms of defence associated with them has thus far been challenging. Advances in our understanding of responses to inconspicuous stimuli, such as those induced by microbial symbionts in herbivore secretions and mechanical stimulation caused by insects, have illuminated the intricacies of herbivory. Here, we provide a synthesis of the interacting impacts of herbivory on plants and the consequential complexities associated with uncoupling defence responses. We propose that simulated herbivory should be used to complement true herbivory to decipher the mechanisms of insect herbivore-induced plant defence responses.

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.

Vegetative plant volatiles influence their environment, and vice versa. We review these dynamic interactions and identify frameworks that can help to explain volatile emission patterns and their biological functions.

Grasses accumulate large amounts of silicon (Si) which is deposited in trichomes, specialised silica cells and cell walls. This may increase leaf toughness and reduce cell rupture, palatability and digestion. Few studies have measured leaf mechanical traits in response to Si, thus the effect of Si on herbivores can be difficult to disentangle from Si-induced changes in leaf surface morphology. We assessed the effects of Si on Brachypodium distachyon mechanical traits (specific leaf area (SLA), thickness, leaf dry matter content (LDMC), relative electrolyte leakage (REL)) and leaf surface morphology (macrohairs, prickle, silica and epidermal cells) and determined the effects of Si on the growth of two generalist insect herbivores (Helicoverpa armigera and Acheta domesticus). Si had no effect on leaf mechanical traits; however, Si changed leaf surface morphology: silica and prickle cells were on average 127% and 36% larger in Si supplemented plants, respectively. Prickle cell density was significantly reduced by Si, while macrohair density remained unchanged. Caterpillars were more negatively affected by Si compared to crickets, possibly due to the latter having a thicker and thus more protective gut lining. Our data show that Si acts as a direct defence against leaf-chewing insects by changing the morphology of specialised defence structures without altering leaf mechanical traits.

Aims: Silicon (Si) uptake and accumulation improves plant resilience to environmental stresses, but most studies examining this functional role of Si have focussed on grasses (Poaceae) and neglected other important plant groups, such as legumes (Fabaceae). Legumes have evolved a symbiotic relationship with nitrogen-fixing bacteria (rhizobia) housed in root nodules. Our study determined the impacts of silicon (Si) supplementation on Medicago truncatula inoculated with Ensifer meliloti rhizobial strains that differed in their capacity for nitrogen fixation: Sm1021 (‘low-efficiency’) or Sm1022 (‘high-efficiency’). Methods: We examined how Si and rhizobial efficacy influence nodule and plant functional traits, including their chemical aspects. These combinations were supplied with or without Si in a glasshouse experiment, where we quantified nodule flavonoids and foliar chemistry (free amino acids, soluble protein, elemental C, N and Si). Results: Si supply increased nodule number per plant, specific nodule flavonoid concentrations, contents of foliar nitrogenous compounds and foliar C, but not foliar Si. We also demonstrated that rhizobial efficacy altered the magnitude of Si effects on certain traits. For example, Si significantly promoted concentrations of foliar N and soluble protein in the plants associated with the ‘low-efficiency’ strain only, and this was not the case with the ‘high-efficiency’ one. Conclusions: Collectively, our study indicates that Si generates positive effects on M. truncatula, particularly when the association with rhizobia is relatively inefficient, and may play a more prominent role in rhizobial functionality than previously thought.