Tansley Review No. 86 Accumulation of phytoalexins: defence mechanism and stimulus response system.

Phytoalexin synthesis is a defence-response- that is characterized by a requirement for a number of distinct elements, all of which must be present for the response to be expressed fully. These same elements: a signal, a cellular receptor, a signal transduction system and a responsive metabolic system, are also used to describe a stimulus-response system. A number of molecular species can function as signal molecules or elicitors of phytoalexin synthesis, including poly- and oligosaccharides, proteins and polypeptides, and fatty acids. Few receptors for elicitors have been identified but those that have been are proteins located on the plasma membrane of the plant. Induction of phytoalexin synthesis involves selective and co-ordinated activation of specific defence response genes, including those encoding the enzymes of phytoalexin synthesis, and these genes constitute the responsive metabolic system. The separate, and distant, locations of the receptor and the responsive genes means that the event in which the signal is perceived by the receptor must be relayed to the genes by means of a second messenger system. Several second messengers are candidates for such a coupling- or signal transduction-system, including udenosine-3',5'-cyclic monophosphate, Ca , diacylglycerol and inositol 1,4,5-trisphosphate, active oxygen species and jasmonic acid. Each has been examined as a possible component of the signal transduction system mediating between the elicitor receptor interaction and the phytoalexin synthesis it induces. Analysis of the signalling events is made complex by the simultaneous solicitation by the invading micro-organism of several defence responses, each of which might involve elements of a different signal system. The same complexity is evident which the role of phytoalexin accumulation in resistance is analysed. Evaluation of the contribution made by phytoalexin accumulation towards resistance has been attempted by the use of various inhibitors and enhancers of the process. Transgenic and mutant plants with specific alterations in one or more ot those elements necessary for the plant to respond to the signals for phytoalexin synthesis and other defence responses, are beginning to aid resolution of the complex pattern ot signalling events and the respective roles of the inducible defence mechanisms in resistance. CONTENTS Summary 1 I. Introduction 2 II. Chemistry of phytoalexins 3 III. Phytoalexin accumulation as a determinant of resistance 6 IV. Elicitation of phytoalexin accumulation 11 References 34.