Phytohormones in Fall Armyworm Saliva Modulate Defense Responses in Plants

Insect herbivory induces plant defense responses that are often modulated by components in insect saliva, oral secretions or regurgitant, frass, or oviposition fluids. These secretions contain proteins and small molecules that act as elicitors or effectors of plant defenses. Several non-protein elicitors have been identified from insect oral secretions, whereas studies of insect saliva have focused mainly on protein identification. Yet, insect saliva may also contain non-protein molecules that could activate defense responses in plants. The goal of this study was to identify non-protein plant defense elicitors present in insect saliva. We used the fall armyworm (FAW), Spodoptera frugiperda and its host plants tomato, maize, and rice as a model system. We tested the effect of protein-digested saliva or non-protein components on herbivore-induced defense responses in maize, rice and tomato. We identified phytohormones in FAW saliva using high performance liquid chromatography coupled with mass spectrometry. The results of this study show that non-protein components in FAW saliva modulated defense responses in different plant species. The saliva of this insect contains benzoic acid, and the phytohormones jasmonic acid, salicylic acid, and abscisic acid at concentrations of <5 ng per μl of saliva. Plant treatment with similar phytohormone quantities detected in FAW saliva upregulated the expression of a maize proteinase inhibitor gene in maize, and down-regulated late herbivore-induced defenses in tomato plants. We conclude that FAW saliva is a complex fluid that, in addition to known enzymatic plant defense elicitors, contains phytohormones and other small molecules.