Cyclization enzymes in the biosynthesis of monoterpenes, sesquiterpenes, and diterpenes.doc

Cyclization enzymes in the biosynthesis of monoterpenes, sesquiterpenes, and diterpenes Cyclization Enzymes in the Biosynthesis of Monoterpenes,Sesquiterpenes,and Diterpenes Edward M.Davis · Rodney Croteau Institute ofBiological Chemistry,Washington State University,Pullman,WA 99164–6340,USAE-mail:croteau@ Terpene synthases catalyze the first committed steps in the biosynthesis ofmonoterpenes,ses-quiterpenes,and diterpenes.An overview is presented ofthe enzymology and mechanism ofthese terpene synthases,and their molecular cloning,expression,and sequence analysis.Detailed structural and functional evaluation offour representative monoterpene,sesquiter-pene,and diterpene synthases is also presented. Keywords.Terpene,Monoterpene,Sesquiterpene,Diterpene,Cyclase,Geranyl diphosphate,Farnesyl diphosphate,Geranylgeranyl diphosphate 1 2 3Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54Prenyl Transferases and Terpenoid Synthases . . . . . . . . . . . . . .58Monoterpene Synthases . . . . . . . . . . . . . . . . . . . . . . . . . .59 3.1Mechanism and Stereochemistry . . . . . . . . . . . . . . . . . . . . .61 3.2Limonene Synthase . . . . . . . . . . . . . . . . . . . . . . . . . . . . .664Sesquiterpene Synthases . . . . . . . . . . . . . . . . . . . . . . . . .72 4.1G-Humulene Synthase and d-Selinene Synthase .