Cytochrome c oxidase: evolution of control via nuclear subunit addition.

Cytochrome c oxidase: evolution of control via nuclear subunit addition.
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DOI:
10.1016/j.bbabio.2011.07.007
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发表时间:
2012-04
影响因子:
4.3
通讯作者:
Grossman, Lawrence I.
Grossman, Lawrence I.
中科院分区:
生物学2区
文献类型:
--
作者:
Pierron, Denis;Wildman, Derek E.;Huettemann, Maik;Markondapatnaikuni, Gopi Chand;Aras, Siddhesh;Grossman, Lawrence I.

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根据理论,现有的真核细胞起源于两个自由生活细胞之间的有益结合。由于这种内共生事件,前真核细胞获得了氧化磷酸化 (OXPHOS),产生的 ATP 是糖酵解的 15 倍以上。由于细胞 ATP 需求波动,而 OXPHOS 需要并产生对真核细胞有毒的实体,如 ROS 或 NADH,因此我们认为内共生的成功在很大程度上取决于内共生体 OXPHOS 的调节。多项研究表明细胞色素 C 氧化酶是 OXPHOS 的关键调节因子;例如,COX 是哺乳动物 OXPHOS 与核编码亚基的已知组织特异性异构体的唯一复合物。我们在这里讨论有关核编码亚基起源以及组织和细胞环境(例如缺氧)促进的不同同工酶出现的当前知识。我们还回顾了最近关于脊椎动物(尤其是灵长类谱系)中 COX 选择性压力作用的证据,并讨论了核基因组和线粒体基因组之间共同进化的独特模式。最后,尽管核编码亚基的添加是真核 COX 进化中的一个重大事件,但这并不会导致更有效的 COX 的出现,正如从人类中心主义的角度所预期的那样,对于拥有大大脑和肌肉的“高等”生物体来说。这些亚基的主要功能似乎“仅”控制线粒体亚基的活性。我们认为这种控制功能是进化的一个尚未被充分认识的关键点。此外,调节能量供应的重要性可能导致在类似于“驯化场景”的过程中添加由细胞核编码的亚基,使得宿主倾向于越来越严格地控制由mtDNA编码的亚基执行的COX的祖先活动。本文是题为“呼吸氧化酶”的特刊的一部分。
According to theory, present eukaryotic cells originated from a beneficial association between two free-living cells. Due to this endosymbiotic event the pre-eukaryotic cell gained access to oxidative phosphorylation (OXPHOS), which produces more than 15 times as much ATP as glycolysis. Because cellular ATP needs fluctuate and OXPHOS both requires and produces entities that can be toxic for eukaryotic cells such as ROS or NADH, we propose that the success of endosymbiosis has largely depended on the regulation of endosymbiont OXPHOS. Several studies have presented cytochrome c oxidase as a key regulator of OXPHOS; for example, COX is the only complex of mammalian OXPHOS with known tissue-specific isoforms of nuclear encoded subunits. We here discuss current knowledge about the origin of nuclear encoded subunits and the appearance of different isozymes promoted by tissue and cellular environments such as hypoxia. We also review evidence for recent selective pressure acting on COX among vertebrates, particularly in primate lineages, and discuss the unique pattern of co-evolution between the nuclear and mitochondrial genomes. Finally, even though the addition of nuclear encoded subunits was a major event in eukaryotic COX evolution, this does not lead to emergence of a more efficient COX, as might be expected from an anthropocentric point of view, for the “higher” organism possessing large brains and muscles. The main function of these subunits appears to be “only” to control the activity of the mitochondrial subunits. We propose that this control function is an as yet underappreciated key point of evolution. Moreover, the importance of regulating energy supply may have caused the addition of subunits encoded by the nucleus in a process comparable to a “domestication scenario” such that the host tends to control more and more tightly the ancestral activity of COX performed by the mtDNA encoded subunits. This article is part of a Special Issue entitled: Respiratory Oxidases.
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