Systemin/Jasmonate-Mediated Systemic Defense Signaling in Tomato

Wound-inducible proteinase inhibitors (PIs) in tomato plants provide a useful model system to elucidate the signal transduction pathways that regulate systemic defense response. Among the proposed intercellular signals for wound-induced PIs expression are the peptide systemin and the oxylipin-derived phytohormone jasmonic acid (JA). An increasing body of evidence indicates that systemin and JA work in the same signaling pathway to activate the expression of PIs and other defense-related genes. However, relatively less is known about how these signals interact to promote cell-to-cell communication over long distances. Genetic analysis of the systemin/JA signaling pathway in tomato plants provides a unique opportunity to study, in a single experimental system, the mechanism by which peptide and oxylipin signals interact to coordinate systemic expression of defense-related genes. Previously, it has been proposed that systemin is the long-distance mobile signal for defense gene expression. Recently, grafting experiments with tomato mutants defective in JA biosynthesis and signaling provide new evidence that JA, rather than systemin, functions as the systemic wound signal, and that the biosynthesis of JA is regulated by the peptide systemin. Further understanding of the systemin/JA signaling pathway promises to provide new insights into the basic mechanisms governing plant defense to biotic stress.