Philippe H, Forterre P
Phylogénie et Evolution Moléculaires (UPRESA 8080 CNRS), Bâtiment 444, Université Paris-Sud, 91405 Orsay-Cedex, France.
J Mol Evol. 1999 Oct;49(4):509-23. doi: 10.1007/pl00006573.
Several composite universal trees connected by an ancestral gene duplication have been used to root the universal tree of life. In all cases, this root turned out to be in the eubacterial branch. However, the validity of results obtained from comparative sequence analysis has recently been questioned, in particular, in the case of ancient phylogenies. For example, it has been shown that several eukaryotic groups are misplaced in ribosomal RNA or elongation factor trees because of unequal rates of evolution and mutational saturation. Furthermore, the addition of new sequences to data sets has often turned apparently reasonable phylogenies into confused ones. We have thus revisited all composite protein trees that have been used to root the universal tree of life up to now (elongation factors, ATPases, tRNA synthetases, carbamoyl phosphate synthetases, signal recognition particle proteins) with updated data sets. In general, the two prokaryotic domains were not monophyletic with several aberrant groupings at different levels of the tree. Furthermore, the respective phylogenies contradicted each others, so that various ad hoc scenarios (paralogy or lateral gene transfer) must be proposed in order to obtain the traditional Archaebacteria-Eukaryota sisterhood. More importantly, all of the markers are heavily saturated with respect to amino acid substitutions. As phylogenies inferred from saturated data sets are extremely sensitive to differences in evolutionary rates, present phylogenies used to root the universal tree of life could be biased by the phenomenon of long branch attraction. Since the eubacterial branch was always the longest one, the eubacterial rooting could be explained by an attraction between this branch and the long branch of the outgroup. Finally, we suggested that an eukaryotic rooting could be a more fruitful working hypothesis, as it provides, for example, a simple explanation to the high genetic similarity of Archaebacteria and Eubacteria inferred from complete genome analysis.
通过祖先基因复制连接的几个复合通用树已被用于确定生命通用树的根。在所有情况下,这个根都位于真细菌分支中。然而,最近从比较序列分析获得的结果的有效性受到了质疑,特别是在古老系统发育的情况下。例如,已经表明,由于进化速率不平等和突变饱和,几个真核生物类群在核糖体RNA或延伸因子树中的位置错误。此外,将新序列添加到数据集中常常会使明显合理的系统发育变得混乱。因此,我们用更新的数据集重新审视了迄今为止用于确定生命通用树根的所有复合蛋白质树(延伸因子、ATP酶、tRNA合成酶、氨甲酰磷酸合成酶、信号识别颗粒蛋白)。一般来说,两个原核生物域不是单系的,在树的不同层次上有几个异常的分组。此外,各自的系统发育相互矛盾,因此必须提出各种特别的设想(旁系同源或横向基因转移)才能获得传统的古细菌-真核生物姐妹关系。更重要的是,所有这些标记在氨基酸替换方面都严重饱和。由于从饱和数据集中推断出的系统发育对进化速率的差异极其敏感,目前用于确定生命通用树根的系统发育可能会受到长枝吸引现象的影响而产生偏差。由于真细菌分支总是最长的分支,真细菌生根可以用该分支与外类群的长分支之间的吸引来解释。最后,我们认为真核生物生根可能是一个更有成效的工作假设,因为例如,它为从全基因组分析推断出的古细菌和真细菌的高遗传相似性提供了一个简单的解释。