Rice defense responses are induced upon leaf rolling by an insect herbivore.

Plant defense against herbivores begins with perception. The earlier plant detects the harm, the greater plant will benefit in its arm race with the herbivore. Before feeding, the larvae of the rice pest Cnaphalocrocis medinalis, initially spin silk and fold up a leaf. Rice can detect and protect itself against C. medinalis feeding. However, whether rice could perceive C. medinalis leaf rolling behavior is currently unknown. Here, we evaluated the role of leaf rolling by C. medinalis and artificial leaf rolling in rice plant defense and its indirect effect on two important C. medinalis parasitoids (Itoplectis naranyae and Apanteles sp.) through a combination of volatile profiling, gene-transcriptional and phytohormonal profiling. Natural leaf rolling by C. medinalis resulted in an increased attraction of I. naranyae when compared to the undamaged plant after 12 h. Volatile analysis revealed that six out of a total 22 components significantly increased in the headspace of C. medinalis rolled plant when compared to undamaged plant. Principal component analysis of these components revealed similarities in the headspace of undamaged plant and artificially rolled plant while the headspace volatiles of C. medinalis rolled plant deferred significantly. Leaf rolling and feeding by C. medinalis up-regulated the plant transcriptome and a series of jasmonic acid (JA) and salicylic acid (SA) related genes. While feeding significantly increased JA level after 12 to 36 h, rolling significantly increased SA level after 2 to 12 h. Compared to artificial rolling, natural rolling significantly increased JA level after 36 h and SA level after 2 and 12 h. Our findings suggest that natural leaf rolling by C. medinalis can be perceived by rice plant. The detection of this behavior may serve as an early warning signal in favor of the rice plant defenses against C. medinalis.