SYSTEMIN: A Polypeptide Signal for Plant Defensive Genes

Damage to leaves of several plant species by herbivores or by other mechanical wounding induces defense gene activation throughout the plants within hours. An 18-amino acid polypeptide, called systemin, has been isolated from tomato leaves that is a powerful inducer of over 15 defensive genes when supplied to the tomato plants at levels of fmol/plant. Systemin is readily transported from wound sites and is considered to be the primary systemic signal. The polypeptide is processed from a 200-amino acid precursor called prosystemin, analogous to polypeptide hormones in animals. However, the plant prohormone does not possess typical dibasic cleavage sites, nor does it contain a signal sequence or any typical membrane-spanning regions. The signal transduction pathway that mediates systemin signaling involves linolenic acid release from membranes and subsequent conversion to jasmonic acid, a potent activator of defense gene transcription. The pathway exhibits analogies to arachidonic acid/prostaglandin signaling in animals that leads to inflammatory and acute phase responses.