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引用次数: 84

摘要

古细菌是一组在最高层次上区别于其他所有生物的生物。它们与其他细菌,即真正的细菌(真细菌)的关系并不比它们与真核细胞的关系更密切。古细菌,真细菌和真核细胞(某些方面)各自代表不同的主要血统。古细菌包括一个小而多样的表型集合。虽然它们具有某些独特的共同表型特征,但很难在这些特征的基础上对它们进行令人信服的分组。当时所需要的(也是所有可靠和令人信服的系统发育测量所需要的)是对家谱关系的基因型中性测量。大分子是计时的,因为其他东西是恒定的,它们序列的变化标志着时间(以随机的,而不是节奏的方式)。因此,比较序列分析是测定家谱关系的有力手段。使用这种(中性的)基因型测量,可以可靠地确定较高水平的系统发育顺序,并且可以将古细菌识别为它们所处的初级王国。虽然现在下结论还为时过早,但似乎古细菌在大多数(如果不是全部的话)分子过程的重要细节上与其他两大类群不同。这与“原核生物”在分子水平上与真核生物不同的方式是一样的。这似乎是不合理的直觉,这三个主要王国之间的这种程度的差异,可以解释为一个表型经历了广泛的进化,成为其他的。更确切地说,似乎所有的物种都有一个共同的祖先,这个祖先拥有更初级、更少细节的表型,因此每个物种都分别进化出了它们不同的细节。普遍的共同祖先应称为祖;它是一种仍处于基因型和表型之间进化联系的阵痛中的生物体。重要的问题是为什么会有一个普遍而独特的祖先。古细菌的存在为真核细胞的起源研究提供了新的有力视角。因此,我们对真核生物起源的概念将进行修订。真核生物起源的传统内共生模型不再是一个充分的解释。真核细胞的主要特征——那些在分子水平上区分真核细胞的特征——早在产生线粒体和叶绿体的内共生现象之前就已经进化出来了。真核细胞的性质可能与祖先的子代的性质有关。古细菌提供的关于进化的问题和答案,应该会在很大程度上重新点燃生物学家对进化问题的兴趣,并有希望在某种程度上把生物学从目前的技术冒险主义道路上转移开来。
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Archaebacteria and Cellular Origins: An Overview

Archaebacteria are a group of organisms distinct from all others at the highest level. They are no more related to other bacteria, i.e., the true bacteria (eubacteria) than they are to eucaryotic cells. Archaebacteria, eubacteria and (some aspect of) the eucaryotic cell each represent separate primary lines of descent.

The archaebacteria comprise a small but diverse collection of phenotypes. While they have certain unique common phenotypic characteristics, it would have been difficult to group them convincingly on the basis of these. What was needed (and what is needed in all reliable and convincing phylogenetic measurement) is a genotypic neutral measure of genealogical relationships. Macromolecules are chronometric, in that other things being constant, changes in their sequences mark time (in a stochastic, not a metronomic way). Comparative sequence analysis is therefore a powerful measure of genealogical relationships. Using such (neutral) genotypic measures, phylogenetic ordering at the higher levels can be reliably determined — and archaebacteria can be recognized for the primary kingdom that they are.

Although it is too early to generalize with certainty, it seems that the archaebacteria differ from the other two major groups in significant details of most, if not all, molecular processes. [This is the same way in which “procaryotes” were known to differ from eucaryotes at the mloecular level.] It does not seem reasonable intuitively that this extent of difference among the three primary kingdoms can be accounted for by one of the phenotypes undergoing an extensive evolution to become the others. Rather, it seems that all lines have shared a common ancestor that possessed a more rudimentary, less detailed, phenotype, and so each has evolved separately the details in which they differ. The universal common ancestor should be called a progenote; it is an organism still in the throes of evolving the link between genotype and phenotype. The important question is why there is a universal, unique ancestor.

The existence of archaebacteria provides a new and powerful perspective on the origin of the eucaryotic cell. As a result, our concept of eucaryotic origins will undergo revision. The conventional endosymbiotic model for eucaryotic origins is no longer a sufficient explanation. The main characteristics of the eucaryotic cell — those that distinguish it at the molecular level — were evolved long before the endosymbioses that led to he mitochon- drion and the chloroplast. The nature of the eucaryotic cell is perhaps related to the nature of the ancestral progenote.

The evolutionary questions and answers offered by archaebacteria should go far to rekindle the biologist's flagging interest in evolutionary matters, and hopefully divert biology to some extent from its present course of technological adventurism.

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