Plant volatiles as regulators of plant defense and herbivore immunity: molecular mechanisms and unanswered questions.

Plants release distinct blends of herbivore-induced plant volatiles (HIPVs) upon herbivore attack. HIPVs have long been known to influence the behavior of herbivores and natural enemies. In addition, HIPVs can act as physiological regulators that induce or prime plant defenses. Recent work indicates that the regulatory capacity of HIPVs may extend to herbivore immunity: herbivores that are exposed to HIPVs can become more resistant or susceptible to parasitoids and pathogens. While the mechanisms of HIPV-mediated plant defense regulation are being unraveled, the mechanisms underlying the regulation of herbivore immunity are unclear. Evidence so far suggests a high degree of context dependency. Here, we review the mechanisms by which HIPVs regulate plant defense and herbivore immunity. We address major gaps of knowledge and discuss directions for future mechanistic research to facilitate efforts to use the regulatory capacity of HIPVs for the biological control of insect pests.