Cazzolla Gatti Roberto
Bio-Clim-Land Centre, Biological Institute, Tomsk State University, Tomsk, Russia.
Theor Biol Forum. 2018 Jan 1;111(1-2):13-26. doi: 10.19272/201811402002.
In 1967 Lynn (Sagan) Margulis proposed that mitochondria, photosynthetic plastids and cilia were acquired prokaryotes and evolved symbiotically to form anaerobic bacteria, photosynthetic bacteria and eventually algae. Although most of this theory is well-accepted now, the hypothesis that endosymbiotic spirochaetes developed into eukaryotic flagella and cilia, and the following proposals of an endosymbiotic origin of other eukaryotic organelles such as peroxisomes, glyoxysomes, etc. have not received much acceptance, since evidence suggests they lack a genome and do not show ultrastructural similarities to bacteria or archaea. Nevertheless, the idea that over millennia mitochondria, plastids, prokaryotic and eukaryotic cells and even flagella and peroxisomes, as either primary or secondary endosymbionts, transferred some or all of their own DNA to the host cells nucleus through a process called «endogenosymbiosis» (i.e. a symbiotic gene transfer, such as the internalisation of the endosymbionts DNA with lateral transfer) has been recently suggested. This endogenosymbiosis could take place during the evolutionary transition from the symbiotic interacting community, invoked by Margulis, to a fully-integrated (either prokaryotic or eukaryotic) cell. This process could explain the missing evidence of the presence of DNA in flagella and peroxisomes whose ancestor endosymbionts, during the long endogenosymbiotic evolution, could have transferred their whole genome to the host cell that subsequently integrated it in its own genome, directly controlling its expression. Furthermore, the endogenosymbiosis hypothesis could be the explanation of the transition between an RNA to a DNA world and of some cases of true sympatric evolution of species, apparently inexplicable by the canonical speciation processes. Here, after an introduction to the theoretical basis of endogenosymbiosis and a discussion of the empirical confirming evidence, I show a graphical summary of the integration between this and the former endosymbiosis theories. The Serial Endosymbiosis Theory and the Secondary Endosymbiosis are merged with the Endogenosymbiosis Theory in a Unified Symbiogenetic Theory (UST).
1967年,林恩·(萨根)·马古利斯提出,线粒体、光合质体和纤毛是被获取的原核生物,并通过共生进化形成了厌氧细菌、光合细菌,最终形成了藻类。尽管该理论的大部分内容如今已被广泛接受,但关于内共生螺旋体演变成真核生物鞭毛和纤毛的假说,以及随后关于其他真核细胞器(如过氧化物酶体、乙醛酸循环体等)起源于内共生的提议,并未得到太多认可,因为有证据表明它们缺乏基因组,且在超微结构上与细菌或古菌没有相似之处。然而,最近有人提出,在数千年的时间里,线粒体、质体、原核细胞和真核细胞,甚至鞭毛和过氧化物酶体,作为初级或次级内共生体,通过一个名为“内共生基因转移”(即内共生体的DNA通过横向转移内化)的过程,将它们自身的部分或全部DNA转移到了宿主细胞的细胞核中。这种内共生基因转移可能发生在从马古利斯提出的共生相互作用群落向完全整合的(原核或真核)细胞的进化转变过程中。这一过程可以解释鞭毛和过氧化物酶体中DNA存在证据缺失的现象,其祖先内共生体在漫长的内共生基因转移进化过程中,可能已将其整个基因组转移到了宿主细胞中,宿主细胞随后将其整合到自身基因组中,并直接控制其表达。此外,内共生基因转移假说可以解释从RNA世界到DNA世界的转变,以及某些物种真正的同域进化案例,而这些案例用经典的物种形成过程显然无法解释。在此,在介绍内共生基因转移的理论基础并讨论实证证据之后,我展示了这一理论与之前内共生理论整合的图形总结。串联内共生理论和次级内共生理论与内共生基因转移理论合并,形成了统一共生起源理论(UST)。