Yakhnin Alexander V
Department of Biochemistry and Molecular Biology, The Pennsylvania State University, University Park, PA 16802, USA.
Orig Life Evol Biosph. 2007 Dec;37(6):523-36. doi: 10.1007/s11084-007-9108-z. Epub 2007 Sep 20.
The origin of protein synthesis is one of the major riddles of molecular biology. It was proposed a decade ago that the ribosomal RNA evolved from an earlier RNA-replisome (a ribozyme fulfilling RNA replication) while transfer RNA (tRNA) evolved from a genomic replication origin. Applying these hypotheses, I suggest that protein synthesis arose for the purpose of segregating copy and template RNA during replication through the conventional formation of a complementary strand. Nascent RNA was scanned in 5' to 3' direction following the progress of replication. The base pairing of several tRNA-like molecules with nascent RNA released the replication intermediates trapped in duplex. Synthesis of random peptides evolved to fuel the turnover of tRNAs. Then the combination of replication-coupled peptide formation and the independent development of amino acid-specific tRNA aminoacylation resulted in template-based protein synthesis. Therefore, the positioning of tRNAs adjacent to each other developed for the purpose of replication rather than peptide synthesis. This hypothesis does not include either selection for useful peptides or specific recognition of amino acids at the initial evolution of translation. It does, however, explain a number of features of modern translation apparatus, such as the relative flexibility of genetic code, the number of proteins shared by the transcription and translation machines, the universal participation of an RNA subunit in co-translational protein secretion, 'unscheduled translation', and factor-independent translocation. Assistance of original ribosomes in keeping apart the nascent transcript from its template is still widely explored by modern bacteria and perhaps by other domains of life.
蛋白质合成的起源是分子生物学的主要谜团之一。十年前有人提出,核糖体RNA由早期的RNA复制体(一种负责RNA复制的核酶)进化而来,而转运RNA(tRNA)则从基因组复制起点进化而来。应用这些假说,我认为蛋白质合成的出现是为了在复制过程中通过互补链的常规形成来分离复制RNA和模板RNA。随着复制的进行,新生RNA从5'端到3'端被扫描。几个类似tRNA的分子与新生RNA的碱基配对释放了被困在双链体中的复制中间体。随机肽的合成逐渐发展以推动tRNA的周转。然后,复制偶联的肽形成与氨基酸特异性tRNA氨酰化的独立发展相结合,导致了基于模板的蛋白质合成。因此,tRNA彼此相邻的定位是为了复制而非肽合成而发展的。这一假说在翻译最初的进化过程中既不包括对有用肽的选择,也不包括对氨基酸的特异性识别。然而,它确实解释了现代翻译装置的一些特征,例如遗传密码的相对灵活性、转录和翻译机器共有的蛋白质数量、RNA亚基在共翻译蛋白质分泌中的普遍参与、“非计划翻译”以及因子非依赖性易位。现代细菌以及或许其他生命域仍在广泛探索原始核糖体在将新生转录本与其模板分开方面的作用。