Embley T Martin, van der Giezen Mark, Horner David S, Dyal Patricia L, Foster Peter
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK.
Philos Trans R Soc Lond B Biol Sci. 2003 Jan 29;358(1429):191-201; discussion 201-2. doi: 10.1098/rstb.2002.1190.
Published data suggest that hydrogenosomes, organelles found in diverse anaerobic eukaryotes that make energy and hydrogen, were once mitochondria. As hydrogenosomes generally lack a genome, the conversion is probably one way. The sources of the key hydrogenosomal enzymes, pyruvate : ferredoxin oxidoreductase (PFO) and hydrogenase, are not resolved by current phylogenetic analyses, but it is likely that both were present at an early stage of eukaryotic evolution. Once thought to be restricted to a few unusual anaerobic eukaryotes, the proteins are intimately integrated into the fabric of diverse eukaryotic cells, where they are targeted to different cell compartments, and not just hydrogenosomes. There is no evidence supporting the view that PFO and hydrogenase originated from the mitochondrial endosymbiont, as posited by the hydrogen hypothesis for eukaryogenesis. Other organelles derived from mitochondria have now been described in anaerobic and parasitic microbial eukaryotes, including species that were once thought to have diverged before the mitochondrial symbiosis. It thus seems possible that all eukaryotes may eventually be shown to contain an organelle of mitochondrial ancestry, to which different types of biochemistry can be targeted. It remains to be seen if, despite their obvious differences, this family of organelles shares a common function of importance for the eukaryotic cell, other than energy production, that might provide the underlying selection pressure for organelle retention.
已发表的数据表明,氢化酶体(存在于多种能产生能量和氢气的厌氧真核生物中的细胞器)曾经是线粒体。由于氢化酶体通常缺乏基因组,这种转变可能是单向的。目前的系统发育分析尚未确定关键氢化酶体酶——丙酮酸:铁氧化还原蛋白氧化还原酶(PFO)和氢化酶的来源,但它们很可能在真核生物进化的早期阶段就已存在。这些蛋白质曾经被认为只存在于少数不寻常的厌氧真核生物中,现在它们紧密地整合到各种真核细胞的结构中,并被靶向到不同的细胞区室,而不仅仅是氢化酶体。没有证据支持真核生物起源的氢化假说所提出的观点,即PFO和氢化酶起源于线粒体共生体。现在已经在厌氧和寄生的微生物真核生物中描述了其他源自线粒体的细胞器,包括一些曾经被认为在线粒体共生之前就已经分化的物种。因此,似乎所有真核生物最终都可能被证明含有一种线粒体祖先的细胞器,不同类型的生物化学过程可以靶向该细胞器。尽管它们存在明显差异,但这个细胞器家族是否除了能量产生之外,还共享对真核细胞至关重要的共同功能,从而可能为细胞器的保留提供潜在的选择压力,仍有待观察。