Plant Biochemistry Laboratory, Department of Plant and Environmental Sciences, University of Copenhagen, Thorvaldsensvej 40, Frederiksberg C, 1871 Copenhagen, Denmark.
Philos Trans R Soc Lond B Biol Sci. 2013 Jan 6;368(1612):20120426. doi: 10.1098/rstb.2012.0426. Print 2013 Feb 19.
The irreversible nature of reactions catalysed by P450s makes these enzymes landmarks in the evolution of plant metabolic pathways. Founding members of P450 families are often associated with general (i.e. primary) metabolic pathways, restricted to single copy or very few representatives, indicative of purifying selection. Recruitment of those and subsequent blooms into multi-member gene families generates genetic raw material for functional diversification, which is an inherent characteristic of specialized (i.e. secondary) metabolism. However, a growing number of highly specialized P450s from not only the CYP71 clan indicate substantial contribution of convergent and divergent evolution to the observed general and specialized metabolite diversity. We will discuss examples of how the genetic and functional diversification of plant P450s drives chemical diversity in light of plant evolution. Even though it is difficult to predict the function or substrate of a P450 based on sequence similarity, grouping with a family or subfamily in phylogenetic trees can indicate association with metabolism of particular classes of compounds. Examples will be given that focus on multi-member gene families of P450s involved in the metabolic routes of four classes of specialized metabolites: cyanogenic glucosides, glucosinolates, mono- to triterpenoids and phenylpropanoids.
P450 催化的反应的不可逆性质使这些酶成为植物代谢途径进化的标志。P450 家族的创始成员通常与一般(即初级)代谢途径相关联,限于单个拷贝或极少数代表,表明存在纯化选择。这些酶的招募以及随后的多成员基因家族的出现,为功能多样化提供了遗传原料,这是专门化(即次生)代谢的固有特征。然而,越来越多的高度专门化的 P450 不仅来自 CYP71 族,还表明趋同和分歧进化对观察到的一般和特殊代谢物多样性有实质性贡献。我们将讨论植物 P450 如何在植物进化的背景下通过遗传和功能多样化来驱动化学多样性的例子。尽管根据序列相似性预测 P450 的功能或底物很困难,但在系统发育树中与家族或亚家族分组可以表明与特定类化合物代谢的关联。将重点介绍参与四类特殊代谢物代谢途径的 P450 多成员基因家族的例子:氰苷葡萄糖苷、硫代葡萄糖苷、单萜类和苯丙素类。