The peritrophic membrane as a target of proteins that play important roles in plant defense and microbial attack

The peritrophic membrane (or peritrophic matrix: PM) is a thin membranous structure that lies along the midgut epithelium in the midgut lumen and consists of chitin and proteins. PM exists between ingested food material and midgut epithelium cells and it is on the frontline of insect-plant and insect-microbe interactions. Therefore, proteins that play major roles in plant defense against herbivorous insects and in microbial attack on insects should penetrate, destroy or modify the PM to accomplish their roles. Recently, it has become clear that some proteins crucial to plant defense or microbial attack have the PM as their primary target. In addition, several plant defense proteins have been reported to affect the PM, although it is still unclear whether the PM is their primary target. This review introduces several of these proteins: fusolin and enhancin, two proteins produced by insect viruses that greatly enhance infection of the viruses by disrupting the PM; the MLX56 family proteins found in mulberry latex as defense proteins against insect herbivores, which modify the PM to a thick structure that inhibits digestive processes; Mir1-CP, a defense cysteine protease from maize that inhibits the growth of insects at very low concentrations and degrades the PM structures; and chitinases and lectins. The importance, necessary characteristics, and modes of action of PM-targeting proteins are then discussed from a strategic point of view, by spotlighting the importance of selective permeability of the PM. Finally, the review discusses the possibility of applying PM-targeting proteins for the control of pest insects.