Plants, for hundreds of millions of years, have been evolving defensive strategies to protect themselves against herbivores and pathogens. As plant species evolved within their ecological niches, they developed their own unique arrays of defensive chemicals against the predators they confronted. Virtually thousands of chemicals are present among living plant species that are thought to be defense-related. The advantages of inducible defensive chemicals to plants is not totally clear, but it can be speculated that many plant species have evolved in very hostile environments in which their nutrients were limited, and the need to conserve energy in diverse ecological systems was essential. The inducible defenses have been of particular interest to biochemists and molecular and cell biologists because of the challenges to understand their complex signaling systems, and because of their potential for genetically regulating defensive genes to improve plant productivity.Defensive chemicals are synthesized in many plant species in response to damage inflicted by attacking herbivores (1, 2). In tomato plants, a signal (or signals) originates at the wound site that is transported throughout the plant where it activates the synthesis of defensive proteins that interfere with the digestive systems of the attacking herbivores (Fig. 1). These inducible defensive proteins have been identified as serine, cysteine and aspartyl proteinase inhibitors (2-4) and polyphenol oxidase (5). These proteins interact with the proteins and proteinases of herbivore guts and adversely affect proteolysis of the ingested food, reducing the availability of essential amino acids and retarding the growth and development of the herbivores (2, 5). The net effect of this process in a natural ecosystem is likely to result in a reduction of damage to the plant, either by killing the predator or by providing a longer period of exposure of surviving herbivores to their natural predators. Research on inducible plant defenses has been primarily concerned with defense against insect predators, but more recently, proteinase inhibitors in sedges and grasses have been found to be a major factor in regulating fluctuating populations of lemmings (6). The initial search for the systemic signal in tomato plants resulted in the finding that oligogalacturonides derived from the plant cell wall were inducers of the defensive proteinase inhibitor genes in excised tomato leaves (7). Subsequently, chitin and chitosan oligomers derived from fungal cell walls were also found to be active inducers (8). However, both classes of oligosaccharides were active only at relatively high concentrations (several hundred micrograms per plant) and did not move to distal leaves when placed on leaf wounds (9). The oligosaccharides are therefore considered to be among the signals produced at sites of pathogen attacks where they help produce localized defensive chemicals. A renewed search for the systemic signal in the soluble components of tomato leaves resulted in the isolation o...