SYSTEMIN: A Polypeptide Signal for Plant Defensive Genes

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Abstract
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.