{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,13]],"date-time":"2026-07-13T15:07:03Z","timestamp":1783955223853,"version":"3.55.0"},"reference-count":46,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["BioSystems"],"published-print":{"date-parts":[[2026,8]]},"DOI":"10.1016\/j.biosystems.2026.105848","type":"journal-article","created":{"date-parts":[[2026,6,7]],"date-time":"2026-06-07T14:27:26Z","timestamp":1780842446000},"page":"105848","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["Domains of life identified through genetic codes"],"prefix":"10.1016","volume":"266","author":[{"given":"Massimo","family":"Di Giulio","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.biosystems.2026.105848_bib1","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2022.104613","article-title":"The genetic code is very close to a global optimum in a model of its origin taking into account both the partition energy of amino acids and their biosynthetic relationships","volume":"214","author":"Caldararo","year":"2022","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib2","doi-asserted-by":"crossref","first-page":"20356","DOI":"10.1073\/pnas.0810647105","article-title":"The archaebacterial origin of eukaryotes","volume":"105","author":"Cox","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105848_bib3","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1016\/0022-2836(68)90392-6","article-title":"The origin of the genetic code","volume":"38","author":"Crick","year":"1968","journal-title":"J. Mol. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib4","series-title":"The God Delusion. Batam Books, United Kingdom","author":"Dawkins","year":"2006"},{"key":"10.1016\/j.biosystems.2026.105848_bib5","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1006\/jtbi.1996.0390","article-title":"On the origin of the genetic code","volume":"187","author":"Di Giulio","year":"1997","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib6","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1006\/jtbi.2001.2271","article-title":"The non-universality of the genetic code: the universal ancestor was a progenote","volume":"209","author":"Di Giulio","year":"2001","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib7","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.biosystems.2004.11.005","article-title":"The origin of the genetic code: theories and their relationships, a review","volume":"80","author":"Di Giulio","year":"2005","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib8","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/1745-6150-3-37","article-title":"An extension of the coevolution theory of the origin of the genetic code","volume":"3","author":"Di Giulio","year":"2008","journal-title":"Biol. Direct"},{"key":"10.1016\/j.biosystems.2026.105848_bib9","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1007\/s00239-010-9407-2","article-title":"The last Universal Common Ancestor (LUCA) and the ancestors of archaea and bacteria were progenotes","volume":"72","author":"Di Giulio","year":"2011","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biosystems.2026.105848_bib10","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/s00239-014-9626-z","article-title":"On how many fundamental kinds of cells are present on Earth: looking for phylogenetic traits that would allow the identification of the primary lines of descent","volume":"78","author":"Di Giulio","year":"2014","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biosystems.2026.105848_bib11","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.jtbi.2018.01.025","article-title":"On Earth, there would be a number of fundamental kinds of primary cells\u2013cellular domains\u2013greater than or equal to four","volume":"443","author":"Di Giulio","year":"2018","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib12","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.jtbi.2018.12.039","article-title":"A qualitative criterion for identifying the root of the tree of life","volume":"464","author":"Di Giulio","year":"2019","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib15","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2023.105014","article-title":"The absence of the evolutionary state of the prokaryote would imply a polyphyletic origin of proteins and that LUCA, the ancestor of bacteria and that of archaea were progenotes","volume":"233","author":"Di Giulio","year":"2023","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib16","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2024.105217","article-title":"Theories of the origin of the genetic code: strong corroboration for the coevolution theory","volume":"239","author":"Di Giulio","year":"2024","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib19","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2026.105787","article-title":"The archaeal genetic code with all UAG codons as pyrrolysine might be an ancestral and not derived trait: the non-monophyletic origin of the genetic code","volume":"263","author":"Di Giulio","year":"2026","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib13","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2020.104330","article-title":"The late appearance of DNA, the nature of the LUCA and ancestors of the domains of life","volume":"202","author":"Di Giulio","year":"2021","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib17","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2024.105382","article-title":"The genetic code is not universal","volume":"247","author":"Di Giulio","year":"2025","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib14","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2021.104449","article-title":"The evolutionary stages of the complexity of biological catalysts mark and clarify the phases of the origin of the genetic code: a model for the origin of the reading frame with codons from proto-mRNAs with different frames","volume":"207","author":"Di Giulio","year":"2021","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib18","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2025.105636","article-title":"How many and which types of primary cells (domains of life) emerged from LUCA, identified employing different genetic codes","volume":"258","author":"Di Giulio","year":"2025","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105848_bib20","doi-asserted-by":"crossref","first-page":"6721","DOI":"10.1073\/pnas.91.15.6721","article-title":"Tempo, mode, the progenote, and the universal root","volume":"91","author":"Doolittle","year":"1994","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105848_bib21","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1101\/SQB.1987.052.01.085","article-title":"The phylogeny of tRNA sequences provides evidence for ambiguity reduction in the origin of the genetic code. Cold spring harbor symp","volume":"52","author":"Fitch","year":"1987","journal-title":"Quant. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib22","first-page":"16","article-title":"The origin of a derived superkingdom: how a gram-positive bacterium crossed the desert to become an archaeon","volume":"vol. 6","author":"Forterre","year":"2011"},{"key":"10.1016\/j.biosystems.2026.105848_bib23","doi-asserted-by":"crossref","first-page":"2197","DOI":"10.1098\/rstb.2009.0034","article-title":"The primary divisions of life: a phylogenomic approach employing composition-heterogeneous methods","volume":"364","author":"Foster","year":"2009","journal-title":"Philos. Trans. R. Soc. B"},{"key":"10.1016\/j.biosystems.2026.105848_bib24","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1126\/science.6771870","article-title":"The phylogeny of prokaryotes","volume":"209","author":"Fox","year":"1980","journal-title":"Science"},{"issue":"3","key":"10.1016\/j.biosystems.2026.105848_bib25","article-title":"Universal paralogs provide a window into evolution before the last universal common ancestor","volume":"6","author":"Goldman","year":"2026","journal-title":"Cell Genom."},{"key":"10.1016\/j.biosystems.2026.105848_bib26","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1146\/annurev.mi.30.100176.002205","article-title":"Enzyme recruitment in evolution of new function","volume":"30","author":"Jensen","year":"1976","journal-title":"Annu. Rev. Microbiol."},{"key":"10.1016\/j.biosystems.2026.105848_bib27","doi-asserted-by":"crossref","first-page":"eadu2404","DOI":"10.1126\/science.adu2404","article-title":"An archaeal genetic code with all TAG codons as pyrrolysine","volume":"390","author":"Kivenson","year":"2025","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105848_bib29","doi-asserted-by":"crossref","first-page":"22","DOI":"10.3390\/life7020022","article-title":"Frozen accident pushing 50: stereochemistry, expansion, and chance in the evolution of the genetic code","volume":"7","author":"Koonin","year":"2017","journal-title":"Life"},{"key":"10.1016\/j.biosystems.2026.105848_bib30","doi-asserted-by":"crossref","first-page":"R1104","DOI":"10.1016\/j.cub.2025.09.071","article-title":"Natural and artificial variations of the standard genetic code","volume":"35","author":"Luke\u0161","year":"2025","journal-title":"Curr. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib31","doi-asserted-by":"crossref","first-page":"1654","DOI":"10.1038\/s41559-024-02461-1","article-title":"The nature of the last universal common ancestor and its impact on the early Earth system","volume":"8","author":"Moody","year":"2024","journal-title":"Nat. Ecol. Evol."},{"key":"10.1016\/j.biosystems.2026.105848_bib32","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/0022-5193(77)90214-4","article-title":"Aminoacyl thiol esters and the origins of genetic specificity","volume":"65","author":"Reanney","year":"1977","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib33","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1007\/s00239-024-10199-4","article-title":"On the nature of the last common ancestor: a story from its translation machinery","volume":"92","author":"Rivas","year":"2024","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biosystems.2026.105848_bib34","series-title":"Evolving Genes and Proteins","article-title":"Degeneracy of the genetic code: extent, nature, and genetic implications","author":"Sonneborn","year":"1965"},{"key":"10.1016\/j.biosystems.2026.105848_bib35","doi-asserted-by":"crossref","DOI":"10.1098\/rsob.170041","article-title":"Domain cell theory supports the independent evolution of the eukarya, bacteria and archaea and the nuclear compartment commonality hypothesis","volume":"7","author":"Staley","year":"2017","journal-title":"Open Biol."},{"key":"10.1016\/j.biosystems.2026.105848_bib36","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1023\/B:BIPH.0000024412.82219.a6","article-title":"The arbitrariness of the genetic code","volume":"19","author":"Stegmann","year":"2004","journal-title":"Biol. Philos."},{"key":"10.1016\/j.biosystems.2026.105848_bib37","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1038\/s41586-024-07677-6","article-title":"The emerging view on the origin and early evolution of eukaryotic cells","volume":"633","author":"Vosseberg","year":"2024","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105848_bib38","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1038\/nature12779","article-title":"An archaeal origin of eukaryotes supports only two primary domains of life","volume":"504","author":"Williams","year":"2013","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105848_bib39","doi-asserted-by":"crossref","first-page":"1546","DOI":"10.1073\/pnas.54.6.1546","article-title":"On the origin of the genetic code","volume":"54","author":"Woese","year":"1965","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105848_bib40","first-page":"221","article-title":"Bacterial evolution","volume":"51","author":"Woese","year":"1987","journal-title":"Microbiol. Res."},{"key":"10.1016\/j.biosystems.2026.105848_bib41","first-page":"789","article-title":"Evolutionary questions: the \u201cprogenote\u201d","volume":"247","author":"Woese","year":"1990","journal-title":"Science 1990"},{"key":"10.1016\/j.biosystems.2026.105848_bib42","doi-asserted-by":"crossref","first-page":"8392","DOI":"10.1073\/pnas.97.15.8392","article-title":"Interpreting the universal phylogenetic tree","volume":"97","author":"Woese","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105848_bib43","doi-asserted-by":"crossref","first-page":"5088","DOI":"10.1073\/pnas.74.11.5088","article-title":"Phylogenetic structure of the prokaryotic domain: the primary kingdoms","volume":"74","author":"Woese","year":"1977","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.biosystems.2026.105848_bib44","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/BF01796132","article-title":"The concept of cellular evolution","volume":"10","author":"Woese","year":"1977","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biosystems.2026.105848_bib45","doi-asserted-by":"crossref","first-page":"12","DOI":"10.3390\/life6010012","article-title":"Coevolution theory of the genetic code at age forty: pathway to translation and synthetic life","volume":"6","author":"Wong","year":"2016","journal-title":"Life"},{"key":"10.1016\/j.biosystems.2026.105848_bib46","doi-asserted-by":"crossref","first-page":"13","DOI":"10.3390\/life7020013","article-title":"The genetic code and RNA-amino acid affinities","volume":"7","author":"Yarus","year":"2017","journal-title":"Life"},{"key":"10.1016\/j.biosystems.2026.105848_bib47","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/S0022-5193(74)80035-4","article-title":"On earlier states of the biochemical system","volume":"44","author":"Ycas","year":"1974","journal-title":"J. Theor. Biol."}],"container-title":["BioSystems"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0303264726001589?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0303264726001589?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,7,13]],"date-time":"2026-07-13T14:28:12Z","timestamp":1783952892000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0303264726001589"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8]]},"references-count":46,"alternative-id":["S0303264726001589"],"URL":"https:\/\/doi.org\/10.1016\/j.biosystems.2026.105848","relation":{},"ISSN":["0303-2647"],"issn-type":[{"value":"0303-2647","type":"print"}],"subject":[],"published":{"date-parts":[[2026,8]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Domains of life identified through genetic codes","name":"articletitle","label":"Article Title"},{"value":"BioSystems","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.biosystems.2026.105848","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 Elsevier B.V. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}],"article-number":"105848"}}