Chitin‐signaling‐dependent responses to insect oral secretions in rice cells propose the involvement of chitooligosaccharides in plant defense against herbivores

Plants recognize molecules related to a variety of biotic stresses through pattern recognition receptors to activate plant immunity. In the interactions between plants and chewing herbivores, such as lepidopteran larvae, oral secretions (OS) are deposited on wounded sites, which results in the elicitation of plant immune responses. The widely conserved receptor-like kinase CHITIN ELICITOR RECEPTOR KINASE 1 (CERK1) has been broadly associated with the recognition of microbial components, such as fungal chitin, but its relevance to herbivory remained unclear. In this study, we used receptor-knockout rice (Oryza sativa) and larvae of the lepidopteran pest Mythimna loreyi to demonstrate that the induction of immune responses triggered by larval OS in rice cells largely depends on CERK1 (OsCERK1). CHITIN ELICITOR-BINDING PROTEIN (CEBiP), an OsCERK1-interacting receptor-like protein that was proposed as the main chitin receptor, also contributed to the responses of rice cells to OS collected from three different lepidopteran species. Furthermore, CEBiP knockout rice seedlings showed lower OS-triggered accumulation of jasmonic acid. These results strongly suggest that the OsCERK1 and CEBiP recognize a particular OS component in chewing lepidopteran herbivores, and point toward the presence of chitooligosaccharides in the OS. Targeted perturbation to chitin recognition, through the use of fungal effector proteins, confirmed the presence of chitooligosaccharides in the OS. Treatments of wounds on rice plants with chitooligosaccharides enhanced a set of immune responses, leading to resistance against an herbivorous insect. Our data show that rice recognizes chitooligosaccharides during larval herbivory to activate resistance, and identifies chitin as a novel herbivore-associated molecular pattern.