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高等植物柠檬酸循环和乙醛酸循环中酶的进化。内共生基因转移的一个案例研究。

Evolution of the enzymes of the citric acid cycle and the glyoxylate cycle of higher plants. A case study of endosymbiotic gene transfer.

作者信息

Schnarrenberger Claus, Martin William

机构信息

Institut für Biologie, Freie Universität Berlin, Germany.

出版信息

Eur J Biochem. 2002 Feb;269(3):868-83. doi: 10.1046/j.0014-2956.2001.02722.x.

Abstract

The citric acid or tricarboxylic acid cycle is a central element of higher-plant carbon metabolism which provides, among other things, electrons for oxidative phosphorylation in the inner mitochondrial membrane, intermediates for amino-acid biosynthesis, and oxaloacetate for gluconeogenesis from succinate derived from fatty acids via the glyoxylate cycle in glyoxysomes. The tricarboxylic acid cycle is a typical mitochondrial pathway and is widespread among alpha-proteobacteria, the group of eubacteria as defined under rRNA systematics from which mitochondria arose. Most of the enzymes of the tricarboxylic acid cycle are encoded in the nucleus in higher eukaryotes, and several have been previously shown to branch with their homologues from alpha-proteobacteria, indicating that the eukaryotic nuclear genes were acquired from the mitochondrial genome during the course of evolution. Here, we investigate the individual evolutionary histories of all of the enzymes of the tricarboxylic acid cycle and the glyoxylate cycle using protein maximum likelihood phylogenies, focusing on the evolutionary origin of the nuclear-encoded proteins in higher plants. The results indicate that about half of the proteins involved in this eukaryotic pathway are most similar to their alpha-proteobacterial homologues, whereas the remainder are most similar to eubacterial, but not specifically alpha-proteobacterial, homologues. A consideration of (a) the process of lateral gene transfer among free-living prokaryotes and (b) the mechanistics of endosymbiotic (symbiont-to-host) gene transfer reveals that it is unrealistic to expect all nuclear genes that were acquired from the alpha-proteobacterial ancestor of mitochondria to branch specifically with their homologues encoded in the genomes of contemporary alpha-proteobacteria. Rather, even if molecular phylogenetics were to work perfectly (which it does not), then some nuclear-encoded proteins that were acquired from the alpha-proteobacterial ancestor of mitochondria should, in phylogenetic trees, branch with homologues that are no longer found in most alpha-proteobacterial genomes, and some should reside on long branches that reveal affinity to eubacterial rather than archaebacterial homologues, but no particular affinity for any specific eubacterial donor.

摘要

柠檬酸循环或三羧酸循环是高等植物碳代谢的核心组成部分,它除了为线粒体内膜的氧化磷酸化提供电子、为氨基酸生物合成提供中间产物,以及为通过乙醛酸循环体中的乙醛酸循环从脂肪酸衍生的琥珀酸进行糖异生提供草酰乙酸外,还发挥着其他作用。三羧酸循环是一条典型的线粒体途径,广泛存在于α-变形菌中,α-变形菌是根据rRNA系统分类定义的真细菌类群,线粒体就起源于此。在高等真核生物中,三羧酸循环的大多数酶都由细胞核编码,并且此前已经证明其中几种酶与其α-变形菌的同源物具有分支关系,这表明真核细胞核基因是在进化过程中从线粒体基因组中获得的。在这里,我们使用蛋白质最大似然系统发育树来研究三羧酸循环和乙醛酸循环中所有酶的个体进化历史,重点关注高等植物中核编码蛋白的进化起源。结果表明,参与这条真核途径的蛋白质中约有一半与其α-变形菌的同源物最为相似,而其余的则与其真细菌(但不是特定的α-变形菌)的同源物最为相似。考虑到(a)自由生活的原核生物之间的横向基因转移过程,以及(b)内共生(共生体到宿主)基因转移的机制,我们发现期望从线粒体的α-变形菌祖先那里获得的所有核基因都能与当代α-变形菌基因组中编码的同源物特异性分支是不现实的。相反,即使分子系统发育学能够完美发挥作用(实际上并非如此),那么从线粒体的α-变形菌祖先那里获得的一些核编码蛋白在系统发育树中也应该与大多数α-变形菌基因组中不再存在的同源物分支,并且有些应该位于显示出与真细菌而非古细菌同源物有亲缘关系但对任何特定真细菌供体没有特殊亲和力的长分支上。

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