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真核细胞产生过程中的细胞分化。

Cellular differentiation in the process of generation of the eukaryotic cell.

作者信息

Nakamura H, Hase A

机构信息

Biological Institute, Faculty of Science Konan University, Kobe, Japan.

出版信息

Orig Life Evol Biosph. 1990;20(6):499-514. doi: 10.1007/BF01808197.

Abstract

Primitive atmosphere of the earth did not contain oxygen gas (O2) when the proto-cells were generated successfully as the result of chemical evolution and then evolved. Therefore, they first had acquired anaerobic energy metabolism, fermentation. The cellular metabolisms have often been formed by reorganizing to combine or recombinate between pre-existing metabolisms and newly born bioreactions. Photosynthetic metabolism in eukaryotic chloroplast consists of an electron-transfer photosystem and a fermentative reductive pentose phosphate cycle. On the other hand, O2-respiration of eukaryotic mitochondrion is made of Embden-Meyerhof (EM) pathway and tricarboxylic acid cycle, which originate from a connection of fermentative metabolisms, and an electron-transfer respiratory chain, which has been derived from the photosystem. These metabolisms already are completed in some evolved prokaryotes, for example the cyanobacterium Chlorogloea fritschii and aerobic photosynthetic bacteria Rhodospirillum rubrum and Erythrobacter sp. Therefore, it can be reasonably presumed that the eukaryotic chloroplast and mitochondrion have once been formed as the result of metabolic (and genetic) differentiations in most evolved cyanobacterium. Symbiotic theory has explained the origin of eukaryotic cell as that in which the mitochondrion and chloroplast have been derived from endosymbionts of aerobic bacterium and cyanobacterium, respectively, and has mentioned as one of the most potent supportive evidences that amino acid sequences of the photosynthetic and O2 -respiratory enzymes show similarities to corresponding prokaryotic enzymes. However, as will be shown in this discussion, many examples have shown currently that prokaryotic sequences of informative molecules are conserved well not only in those of the mitochondrial and chloroplast molecules but also in the nuclear molecules. In fact, the similarities in sequence of informative molecules are preserved well among the organisms not only in phylogenetically close relationships but also under highly selective pressure, that is under a physiological constraint for the species in their habitats. Therefore, the similarities in amino acid sequences of proteins between the prokaryotes and the organelles are not necessarily direct evidence for their phylogenetical closeness: it gives still less evidence for a symbiotic relationship between the prokaryotes and the organelles. The metabolic compartmentalization of the membranes is an important tendency in cellular evolution to guarantee high specificity and rate of the metabolisms. It is suggested from the data that the intracellular membranes are not static but undergo dynamic turnover. Furthermore, these facts strongly support the Membrane Evolution Theory which was proposed by one of the authors in 1975.

摘要

当地球上的原始细胞通过化学进化成功产生并随后进化时,其原始大气中并不含有氧气(O₂)。因此,它们最初获得的是无氧能量代谢,即发酵作用。细胞代谢常常是通过对既有代谢过程与新生生物反应进行组合或重组而形成的。真核叶绿体中的光合代谢由一个电子传递光合系统和一个发酵性还原戊糖磷酸循环组成。另一方面,真核线粒体的有氧呼吸由糖酵解(EM)途径和三羧酸循环构成,这两者源自发酵代谢的关联,还有一个源自光合系统的电子传递呼吸链。这些代谢过程在一些进化的原核生物中就已经完善,比如绿球藻和红色红螺菌以及赤杆菌。因此,可以合理推测真核叶绿体和线粒体曾经是在最进化的蓝细菌中代谢(和遗传)分化的结果。共生理论解释了真核细胞的起源,即线粒体和叶绿体分别源自需氧细菌和蓝细菌的内共生体,并将光合和有氧呼吸酶的氨基酸序列与相应原核酶显示出相似性作为最有力的支持证据之一。然而,正如本次讨论将表明的,目前许多例子显示,信息分子的原核序列不仅在线粒体和叶绿体分子中,而且在核分子中都保存得很好。事实上,信息分子序列的相似性不仅在系统发育关系密切的生物之间,而且在高度选择压力下,即在物种栖息地的生理限制下,在生物之间都保存得很好。因此,原核生物和细胞器之间蛋白质氨基酸序列的相似性不一定是它们系统发育关系密切的直接证据:它更不能作为原核生物和细胞器之间存在共生关系的证据。膜的代谢区室化是细胞进化中的一个重要趋势,以确保代谢的高特异性和速率。数据表明细胞内膜并非静止不变,而是经历动态更新。此外,这些事实有力地支持了作者之一在1975年提出的膜进化理论。

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