In tomato plants, systemic induction of defense genes in response to herbivory or mechanical wounding is regulated by an 18-amino-acid
peptide signal called
systemin. Transgenic plants that overexpress prosystemin, the
systemin precursor, from a 35S::prosystemin (35S::prosys) transgene exhibit constitutive expression of
wound-inducible defense
proteins including
proteinase inhibitors and
polyphenol oxidase. To study further the role of (pro)
systemin in the
wound response pathway, we isolated and characterized mutations that suppress 35S::prosys-mediated phenotypes. Ten recessive, extragenic suppressors were identified. Two of these define new alleles of def-1, a previously identified mutation that blocks both
wound- and
systemin-induced gene expression and renders plants susceptible to herbivory. The remaining mutants defined four loci designated Spr-1, Spr-2, Spr-3, and Spr-4 (for Suppressed in 35S::prosystemin-mediated responses). spr-3 and spr-4 mutants were not significantly affected in their response to either
systemin or mechanical wounding. In contrast, spr-1 and spr-2 plants lacked systemic
wound responses and were insensitive to
systemin. These results confirm the function of (pro)
systemin in the transduction of systemic
wound signals and further establish that wounding,
systemin, and 35S::prosys induce defensive gene expression through a common signaling pathway defined by at least three genes (Def-1, Spr-1, and Spr-2).