植物天然农药除虫菊酯的生物合成和应用研究进展
Biosynthesis and application of pyrethrins: a natural pesticide from plants
（除虫菊酯的配体合成起始于在除虫菊花器官的子房外壁腺体腺毛中，TcCDS催化2分子的DMAPP生成菊醇二磷酸，并在磷酸水解酶TcNudix1、脱氢酶TcADH2和TcALDH1的作用下生成菊酸，以及另外的氧化酶TcCHH和甲基转移酶TcCCMT的参与下形成第二菊酸；醇配体的前体茉莉酮通过茉莉酸的合成途径生成，而下游醇配体的合成由细胞色素P450酶TcJMH和TcPYS催化形成茉莉酮醇和除虫酮醇，瓜菊酮醇的合成途径尚不清楚；两种醇配体和三种酸配体在TcGLIP酶的催化下形成六种化合物。酸配体合成前期在质体中进行，后进入细胞质中进行进一步的氧化；茉莉酸的合成前体在质体中，后进入过氧化物酶体进行进一步反应，参与醇配体合成的细胞色素P450定位于内质网。分别定位于质体膜和过氧化物酶体膜的转运蛋白Jassy和CTS参与茉莉酸的合成，菊醇从质体中运出到细胞质和各配体从腺体腺毛运送到胞间是否需要转运蛋白还需要进一步验证）
(Synthesis of pyrethrin moieties are originated in the ovary trichome, TcCDS catalyzes 2 DMAPP to generate chrysanthemyl diphosphate, which is further catalyzed by phosphatase TcNudix1, dehydrogenases TcADH2 and TcALDH1 to generate chrysanthemic acid, and two additional enzymes oxidase TcCHH and methyltransferase TcCCMT participate the reaction to form pyrethric acid. Alcohol moieties are generated from the jasmonic acid biosynthesis pathway, and the downstream biosynthesis are catalyzed by cytochrome P450 TcJMH and TcPYS for the biosynthesis of jasmolone and pyrethrolone, but the reactions for cinerolone biosynthesis are still unknown. One of two acid moieties and one of three alcohol moieties are condensed by the catalysis of TcGLIP to produce six different pyrethrins. Upstream steps of the acid moiety pathway are located within plastid, and then the intermediates are transferred to cytosol for further oxidation. The biosynthesis of jasmonic acid is in the plastid and peroxisome under the catalysis of the cytochrome P450s localized at endoplasmic reticulum. Two transporter proteins Jassy and CTS involved in the biosynthesis of jasmonic acid are localized at the membrane of plastid and peroxisome, respectively, and more transporters may exist for transferring chrysanthemol from plastid to cytosol and also for transferring moieties from trichome to apoplast, which need further validation)
