An intracellular signaling pathway for activating plant defense genes against attacking herbivores and pathogens is mediated by a lipid-based signal transduction cascade. I n this pathway, linolenic acid (18:3) is proposed to be liberated from cell membranes and is converted to cyclopentanones that are involved in transcriptional regulation of defense genes, analogously to prostaglandin synthesis and function in animals. Levels of 18:3 and linoleic acid in tomato (Lycopersicon esculentum) leaves increased within 1 h when the leaves were wounded with a hemostat across the main vein t o simulate herbivore attacks. The increase correlated with the time course of accumulation of jasmonic acid, a cyciopentanone product of 18:3, that had previously been shown t o increase i n leaves in response both to wounding and to elicitors of plant defense genes. One hour after wounding, at least a 15-fold excess of 18:3 was found over that required to account for the levels of newly synthesized jasmonic acid. The free fatty acids in both control and wounded leaves accounted for less than 0.25% of the total fatty acids. However, the total lipid contents of the leaves remained relatively unchanged up to 8 h after wounding, indicating that extensive loss of lipids did not occur, although a gradual decrease in polar lipids was observed, mainly in monogalactosyl diacylglycerol of chloroplast lipids. The data support a role for lipid release as a key step in the signaling events that activate defense genes in tomato leaves in response t o wounding by attacking herbivores.
In leaves of tomato (Lycopersicon esculentum) plants, severa1defense genes are activated by herbivore attacks or other mechanical wounds that crush the tissues (Ryan, 1992; Schaller and Ryan, 1996). Among the activated genes are those encoding Ser, aspartyl, and Cys proteinase i h b i t o r s and the enzyme polyphenol oxidase. These genes are activated in leaves both nearby and distant from the wound sites (Ryan, 1992; Schaller and Ryan, 1996). The signals released to local cells have been attributed to (a) oligosaccharide fragments of the plant and/or pathogen cell walls (Bishop et al., 1984;Darvill and Albersheim, 1984), (b) 18:3 (Farmer and Ryan, 1992), a major component of plant membranes, and (c)