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The systemic accumulation of both hydrogen peroxide (H202) and proteinase inhibitor proteins in tomato leaves in response to wounding was inhibited by the NADPH oxidase inhibitors diphenylene iodonium (DPI), imidazole, and pyridine. The expression of several defense genes in response to wounding, systemin, oligosaccharides, and methyl jasmonate also was inhibited by DPI. These genes, including those of four proteinase inhibitors and polyphenol oxidase, are expressed within 4 to 12 hr after wounding. However, DPI did not inhibit the wound-inducible expression of genes encoding prosystemin, lipoxygenase, and allene oxide synthase, which are associated with the octadecanoid signaling pathway and are expressed 0.5 to 2 hr after wounding. Accordingly, treatment of plants with the H 202-generating enzyme glucose oxidase plus glucose resulted in the induction of only the later-expressed defensive genes and not the early-expressed signaling-related genes. H202 was cytochemically detected in the cell walls of vascular parenchyma cells and spongy mesophyll cells within 4 hr after wounding of wild-type tomato leaves, but not earlier. The cumulative results suggest that active oxygen species are generated near cell walls of vascular bundle cells by oligogalacturonide fragments produced by wound-inducible polygalacturonase and that the resulting H202 acts as a second messenger for the activation of defense genes in mesophyll cells. These data provide a rationale for the sequential, coordinated, and functional roles of systemin, jasmonic acid, oligogalacturonides, and H202 signals for systemic signaling in tomato plants in response to wounding.
INTRODUCTION
Reactive oxygen species (ROS) are common components of the defense responses of plants against pathogen and herbivore attacks. Inoculation of plant tissues with pathogens or treatment of cell cultures with microbial elicitors causes an oxidative burst characterized by the rapid generation of hydrogen peroxide (H202; reviewed by Low and Merida, 1996; Lamb and Dixon, 1997; Bolwell, 1999). Similarly, ROS are generated in plant tissues in response to wounding (Angelini et al., 1990; Bradley et al., 1992; Olson and Varner, 1993; Felton et al., 1994; Bi and Felton, 1995; Orozco-- Cardenas and Ryan, 1999). Mechanical stimulation of isolated cells (Yahraus et al., 1995; Gus-Mayer et al., 1998) and the treatment of cell suspension cultures with plant cell wall-- derived oligogalacturonic acid (OGA; Legendre et al., 1993; Stennis et al., 1998) also generate H202 accumulation. Wound-- induced H202 accumulation...