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Yet, its origin is still unclear. A structural analysis has shown that the stability of the Watson\u2013Crick base pair at the second position of the anticodon\u2013codon interaction is a critical parameter controlling the extent of non-specific pairings accepted at the third position by the ribosome, a flexibility at the root of degeneracy. Based on recent cryo-EM analyses, the present work shows that residue A1493 of the decoding center provides a significant contribution to the stability of this base pair, revealing that the ribosome is directly involved in the establishment of degeneracy. Building on existing evolutionary models, we show the evidence that the early appearance of A1493 and A1492 established the basis of degeneracy when an elementary kinetic scheme of translation was prevailing. Logical considerations on the expansion of this kinetic scheme indicate that the acquisition of the peptidyl transferase center was the next major evolutionary step, while the induced-fit mechanism, that enables a sharp selection of the tRNAs, necessarily arose later when G530 was acquired by the decoding center.<\/jats:p>","DOI":"10.1093\/nar\/gkac171","type":"journal-article","created":{"date-parts":[[2022,3,23]],"date-time":"2022-03-23T16:10:41Z","timestamp":1648051841000},"page":"4113-4126","source":"Crossref","is-referenced-by-count":18,"title":["Genetic code degeneracy is established by the decoding center of the ribosome"],"prefix":"10.1093","volume":"50","author":[{"given":"Shixin","family":"Ye","sequence":"first","affiliation":[{"name":"INSERM U1195 unit, University of Paris-Saclay, 94276 Le Kremlin Bic\u00eatre, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9887-8123","authenticated-orcid":false,"given":"Jean","family":"Lehmann","sequence":"additional","affiliation":[{"name":"Institute for Integrative Biology of the Cell (I2BC), CEA, CNRS, University of Paris-Saclay, 91198 Gif-sur-Yvette, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2022,3,24]]},"reference":[{"key":"2022042200033328900_B1","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/j.tibs.2003.11.009","article-title":"Historical review: deciphering the genetic code\u2013a personal account","volume":"29","author":"Nirenberg","year":"2004","journal-title":"Trends Biochem. 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