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href="#Inferring_LUCA's_features"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Inferring LUCA's features</span> </div> </a> <button aria-controls="toc-Inferring_LUCA's_features-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Inferring LUCA's features subsection</span> </button> <ul id="toc-Inferring_LUCA's_features-sublist" class="vector-toc-list"> <li id="toc-An_anaerobic_thermophile" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#An_anaerobic_thermophile"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>An anaerobic thermophile</span> </div> </a> <ul id="toc-An_anaerobic_thermophile-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_interpretations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Alternative_interpretations"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Alternative interpretations</span> </div> </a> <ul id="toc-Alternative_interpretations-sublist" class="vector-toc-list"> <li id="toc-Evidence_that_LUCA_was_mesophilic" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Evidence_that_LUCA_was_mesophilic"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.1</span> <span>Evidence that LUCA was mesophilic</span> </div> </a> <ul id="toc-Evidence_that_LUCA_was_mesophilic-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Age" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Age"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Age</span> </div> </a> <ul id="toc-Age-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Root_of_the_tree_of_life" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Root_of_the_tree_of_life"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Root of the tree of life</span> </div> </a> <ul id="toc-Root_of_the_tree_of_life-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-LUCA_and_viruses" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#LUCA_and_viruses"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>LUCA and viruses</span> </div> </a> <ul id="toc-LUCA_and_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Notes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Notes</span> </div> </a> <ul id="toc-Notes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Last universal common ancestor</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 40 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-40" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">40 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Laaste_universele_gemeenskaplike_voorouer" title="Laaste universele gemeenskaplike voorouer – Afrikaans" lang="af" hreflang="af" data-title="Laaste universele gemeenskaplike voorouer" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B3%D9%84%D9%81_%D8%B4%D8%A7%D9%85%D9%84_%D8%A3%D8%AE%D9%8A%D8%B1" title="سلف شامل أخير – Arabic" lang="ar" hreflang="ar" data-title="سلف شامل أخير" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9F%D0%BE%D1%81%D0%BB%D0%B5%D0%B4%D0%B5%D0%BD_%D1%83%D0%BD%D0%B8%D0%B2%D0%B5%D1%80%D1%81%D0%B0%D0%BB%D0%B5%D0%BD_%D0%BE%D0%B1%D1%89_%D0%BF%D1%80%D0%B5%D0%B4%D1%88%D0%B5%D1%81%D1%82%D0%B2%D0%B5%D0%BD%D0%B8%D0%BA" title="Последен универсален общ предшественик – Bulgarian" lang="bg" hreflang="bg" data-title="Последен универсален общ предшественик" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/%C3%9Altim_avantpassat_com%C3%BA_universal" title="Últim avantpassat comú universal – Catalan" lang="ca" hreflang="ca" data-title="Últim avantpassat comú universal" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%AE%D0%BB%D0%B0%D1%88%D0%BA%D0%B8_%D1%83%D0%BD%D0%B8%D0%B2%D0%B5%D1%80%D1%81%D0%B0%D0%BB%D0%BB%C4%83_%D0%BF%C4%95%D1%80%D0%BB%C4%95%D1%85%D0%BB%C4%95_%D0%BD%D0%B5%D1%81%C4%95%D0%BB" title="Юлашки универсаллă пĕрлĕхлĕ несĕл – Chuvash" lang="cv" hreflang="cv" data-title="Юлашки универсаллă пĕрлĕхлĕ несĕл" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target"><span>Чӑвашла</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Posledn%C3%AD_univerz%C3%A1ln%C3%AD_spole%C4%8Dn%C3%BD_p%C5%99edek" title="Poslední univerzální společný předek – Czech" lang="cs" hreflang="cs" data-title="Poslední univerzální společný předek" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Sidste_f%C3%A6lles_forfader" title="Sidste fælles forfader – Danish" lang="da" hreflang="da" data-title="Sidste fælles forfader" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Urvorfahr" title="Urvorfahr – German" lang="de" hreflang="de" data-title="Urvorfahr" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Viimane_%C3%BChine_eellane" title="Viimane ühine eellane – Estonian" lang="et" hreflang="et" data-title="Viimane ühine eellane" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/%C3%9Altimo_antepasado_com%C3%BAn_universal" title="Último antepasado común universal – Spanish" lang="es" hreflang="es" data-title="Último antepasado común universal" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Azken_arbaso_komun_unibertsala" title="Azken arbaso komun unibertsala – Basque" lang="eu" hreflang="eu" data-title="Azken arbaso komun unibertsala" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A2%D8%AE%D8%B1%DB%8C%D9%86_%D9%86%DB%8C%D8%A7%DB%8C_%D9%85%D8%B4%D8%AA%D8%B1%DA%A9_%D8%AC%D9%87%D8%A7%D9%86%DB%8C" title="آخرین نیای مشترک جهانی – Persian" lang="fa" hreflang="fa" data-title="آخرین نیای مشترک جهانی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Dernier_anc%C3%AAtre_commun_universel" title="Dernier ancêtre commun universel – French" lang="fr" hreflang="fr" data-title="Dernier ancêtre commun universel" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/%C3%9Altimo_antepasado_com%C3%BAn_universal" title="Último antepasado común universal – Galician" lang="gl" hreflang="gl" data-title="Último antepasado común universal" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%AA%A8%EB%93%A0_%EC%83%9D%EB%AC%BC%EC%9D%98_%EA%B3%B5%ED%86%B5_%EC%A1%B0%EC%83%81" title="모든 생물의 공통 조상 – Korean" lang="ko" hreflang="ko" data-title="모든 생물의 공통 조상" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8E%D5%A5%D6%80%D5%BB%D5%AB%D5%B6_%D5%B8%D6%82%D5%B6%D5%AB%D5%BE%D5%A5%D6%80%D5%BD%D5%A1%D5%AC_%D5%A8%D5%B6%D5%A4%D5%B0%D5%A1%D5%B6%D5%B8%D6%82%D6%80_%D5%B6%D5%A1%D5%AD%D5%B6%D5%AB" title="Վերջին ունիվերսալ ընդհանուր նախնի – Armenian" lang="hy" hreflang="hy" data-title="Վերջին ունիվերսալ ընդհանուր նախնի" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Leluhur_bersama_terakhir_universal" title="Leluhur bersama terakhir universal – Indonesian" lang="id" hreflang="id" data-title="Leluhur bersama terakhir universal" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/S%C3%AD%C3%B0asti_sameiginlegi_forfa%C3%B0ir_allra_n%C3%BAlifandi_l%C3%ADfvera" title="Síðasti sameiginlegi forfaðir allra núlifandi lífvera – Icelandic" lang="is" hreflang="is" data-title="Síðasti sameiginlegi forfaðir allra núlifandi lífvera" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Ultimo_antenato_comune_universale" title="Ultimo antenato comune universale – Italian" lang="it" hreflang="it" data-title="Ultimo antenato comune universale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%90%D7%91_%D7%94%D7%A7%D7%93%D7%9E%D7%95%D7%9F_%D7%94%D7%9B%D7%95%D7%9C%D7%9C" title="האב הקדמון הכולל – Hebrew" lang="he" hreflang="he" data-title="האב הקדמון הכולל" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A3%E1%83%99%E1%83%90%E1%83%9C%E1%83%90%E1%83%A1%E1%83%99%E1%83%9C%E1%83%94%E1%83%9A%E1%83%98_%E1%83%A3%E1%83%9C%E1%83%98%E1%83%95%E1%83%94%E1%83%A0%E1%83%A1%E1%83%90%E1%83%9A%E1%83%A3%E1%83%A0%E1%83%98_%E1%83%A1%E1%83%90%E1%83%94%E1%83%A0%E1%83%97%E1%83%9D_%E1%83%AC%E1%83%98%E1%83%9C%E1%83%90%E1%83%9E%E1%83%90%E1%83%A0%E1%83%98" title="უკანასკნელი უნივერსალური საერთო წინაპარი – Georgian" lang="ka" hreflang="ka" data-title="უკანასკნელი უნივერსალური საერთო წინაპარი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Ultimus_antecessor_universalis" title="Ultimus antecessor universalis – Latin" lang="la" hreflang="la" data-title="Ultimus antecessor universalis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%9C%E0%B5%80%E0%B4%B5%E0%B4%A8%E0%B5%8D%E0%B4%B1%E0%B5%86_%E0%B4%85%E0%B4%B5%E0%B4%B8%E0%B4%BE%E0%B4%A8_%E0%B4%AA%E0%B5%8A%E0%B4%A4%E0%B5%81_%E0%B4%AA%E0%B5%82%E0%B5%BC%E0%B4%B5%E0%B5%8D%E0%B4%B5%E0%B4%BF%E0%B4%95%E0%B5%BB" title="ജീവന്റെ അവസാന പൊതു പൂർവ്വികൻ – Malayalam" lang="ml" hreflang="ml" data-title="ജീവന്റെ അവസാന പൊതു പൂർവ്വികൻ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Leluhur_universal_terakhir" title="Leluhur universal terakhir – Malay" lang="ms" hreflang="ms" data-title="Leluhur universal terakhir" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/LUCA_(evolutie)" title="LUCA (evolutie) – Dutch" lang="nl" hreflang="nl" data-title="LUCA (evolutie)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%88%D8%B1%D9%88%D8%B3%D8%AA%DB%8C_%D9%86%DA%93%DB%8C%D9%88%D8%A7%D9%84_%D9%85%D8%B4%D8%AA%D8%B1%DA%A9_%D8%AC%D8%AF/%D9%86%DB%8C%DA%A9%D9%87" title="وروستی نړیوال مشترک جد/نیکه – Pashto" lang="ps" hreflang="ps" data-title="وروستی نړیوال مشترک جد/نیکه" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Ostatni_uniwersalny_wsp%C3%B3lny_przodek" title="Ostatni uniwersalny wspólny przodek – Polish" lang="pl" hreflang="pl" data-title="Ostatni uniwersalny wspólny przodek" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/%C3%9Altimo_ancestral_comum_universal" title="Último ancestral comum universal – Portuguese" lang="pt" hreflang="pt" data-title="Último ancestral comum universal" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Ultimul_str%C4%83mo%C8%99_comun_universal" title="Ultimul strămoș comun universal – Romanian" lang="ro" hreflang="ro" data-title="Ultimul strămoș comun universal" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17559452 badge-recommendedarticle mw-list-item" title="recommended article"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BE%D1%81%D0%BB%D0%B5%D0%B4%D0%BD%D0%B8%D0%B9_%D1%83%D0%BD%D0%B8%D0%B2%D0%B5%D1%80%D1%81%D0%B0%D0%BB%D1%8C%D0%BD%D1%8B%D0%B9_%D0%BE%D0%B1%D1%89%D0%B8%D0%B9_%D0%BF%D1%80%D0%B5%D0%B4%D0%BE%D0%BA" title="Последний универсальный общий предок – Russian" lang="ru" hreflang="ru" data-title="Последний универсальный общий предок" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sl 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Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Most recent common ancestor of all current life on Earth</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">"LUCA" redirects here. For other uses, see <a href="/wiki/Luca_(disambiguation)" class="mw-redirect mw-disambig" title="Luca (disambiguation)">Luca</a>.</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Phylogenetic_tree_of_life_1990_LUCA.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Phylogenetic_tree_of_life_1990_LUCA.svg/390px-Phylogenetic_tree_of_life_1990_LUCA.svg.png" decoding="async" width="390" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Phylogenetic_tree_of_life_1990_LUCA.svg/585px-Phylogenetic_tree_of_life_1990_LUCA.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Phylogenetic_tree_of_life_1990_LUCA.svg/780px-Phylogenetic_tree_of_life_1990_LUCA.svg.png 2x" data-file-width="1860" data-file-height="1010" /></a><figcaption><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogenetic tree</a> linking all major groups of living organisms, namely the <a href="/wiki/Bacteria" title="Bacteria">Bacteria</a>, <a href="/wiki/Archaea" title="Archaea">Archaea</a>, and <a href="/wiki/Eukaryote" title="Eukaryote">Eukarya</a>, as proposed by Woese et al 1990,<sup id="cite_ref-Woese_Kandler_Wheelis_1990_1-0" class="reference"><a href="#cite_note-Woese_Kandler_Wheelis_1990-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> with the last universal common ancestor (LUCA) shown at the root</figcaption></figure> <p>The <b>last universal common ancestor</b> (<b>LUCA</b>) is the hypothesized common ancestral <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> from which the <a href="/wiki/Three-domain_system" title="Three-domain system">three domains of life</a>, the <a href="/wiki/Bacteria" title="Bacteria">Bacteria</a>, the <a href="/wiki/Archaea" title="Archaea">Archaea</a>, and the <a href="/wiki/Eukarya" class="mw-redirect" title="Eukarya">Eukarya</a> originated. The cell had a <a href="/wiki/Lipid_bilayer" title="Lipid bilayer">lipid bilayer</a>; it possessed the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> and <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a> which <a href="/wiki/Translation_(biology)" title="Translation (biology)">translated</a> from <a href="/wiki/DNA" title="DNA">DNA</a> or <a href="/wiki/RNA" title="RNA">RNA</a> to <a href="/wiki/Protein" title="Protein">proteins</a>. The LUCA probably existed at latest 3.6 billion years ago, and possibly as early as 4.3 billion years ago<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> or earlier. The nature of this point or stage of divergence remains a topic of research. </p><p>All earlier forms of life preceding this divergence and all extant organisms are generally thought to share <a href="/wiki/Common_ancestry" class="mw-redirect" title="Common ancestry">common ancestry</a>. On the basis of a formal statistical test, this theory of a universal common ancestry (UCA) is supported versus competing multiple-ancestry hypotheses. The <a href="/wiki/First_universal_common_ancestor" title="First universal common ancestor">first universal common ancestor</a> (FUCA) is a hypothetical non-cellular ancestor to LUCA and other now-extinct sister lineages. </p><p>Whether the genesis of <a href="/wiki/Virus" title="Virus">viruses</a> falls before or after the LUCA–as well as the diversity of extant viruses and their hosts–remains a subject of investigation. </p><p>While no fossil evidence of the LUCA exists, the detailed biochemical similarity of all current life (divided into the three domains) makes its existence widely accepted by biochemists. Its characteristics can be inferred from <a href="/wiki/Phylogenetic_bracketing" title="Phylogenetic bracketing">shared features of modern genomes</a>. These genes describe a complex life form with many <a href="/wiki/Co-adapted" class="mw-redirect" title="Co-adapted">co-adapted</a> features, including <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcription</a> and <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> mechanisms to convert information from <a href="/wiki/DNA" title="DNA">DNA</a> to <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> to <a href="/wiki/Protein" title="Protein">proteins</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Historical_background">Historical background</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=1" title="Edit section: Historical background"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">Tree of life (biology)</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Darwin_Tree_1837.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Darwin_Tree_1837.png/170px-Darwin_Tree_1837.png" decoding="async" width="170" height="277" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Darwin_Tree_1837.png/255px-Darwin_Tree_1837.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/10/Darwin_Tree_1837.png/340px-Darwin_Tree_1837.png 2x" data-file-width="1390" data-file-height="2265" /></a><figcaption>A <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a>, like this one from <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>'s notebooks <abbr title="circa">c.</abbr><span style="white-space:nowrap;"> July 1837</span>, implies a single common ancestor at its root (labelled "1").</figcaption></figure> <p>A <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a> directly portrays the idea of <a href="/wiki/Evolution" title="Evolution">evolution</a> by <a href="/wiki/Common_descent" title="Common descent">descent from a single ancestor</a>.<sup id="cite_ref-Gregory_2008_3-0" class="reference"><a href="#cite_note-Gregory_2008-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> An early tree of life was sketched by <a href="/wiki/Jean-Baptiste_Lamarck" title="Jean-Baptiste Lamarck">Jean-Baptiste Lamarck</a> in his <i><a href="/wiki/Philosophie_zoologique" title="Philosophie zoologique">Philosophie zoologique</a></i> in 1809.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> more famously proposed the theory of universal common descent through an evolutionary process in his book <i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i> in 1859: "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed."<sup id="cite_ref-Darwin_1859_6-0" class="reference"><a href="#cite_note-Darwin_1859-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The last sentence of the book begins with a restatement of the hypothesis: </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one ...</p><div class="templatequotecite">— <cite><sup id="cite_ref-Darwin_1859_6-1" class="reference"><a href="#cite_note-Darwin_1859-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p>The term "last universal common ancestor" or "LUCA" was first used in the 1990s for such a primordial organism.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Forterre_1997_pp._764–770_8-0" class="reference"><a href="#cite_note-Forterre_1997_pp._764–770-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Inferring_LUCA's_features"><span id="Inferring_LUCA.27s_features"></span>Inferring LUCA's features</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=2" title="Edit section: Inferring LUCA's features"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="An_anaerobic_thermophile">An anaerobic thermophile</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=3" title="Edit section: An anaerobic thermophile"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Phylogenetic_bracketing" title="Phylogenetic bracketing">Phylogenetic bracketing</a></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tnone center"><div class="thumbinner multiimageinner" style="width:842px;max-width:842px"><div class="trow"><div class="tsingle" style="width:276px;max-width:276px"><div class="thumbimage" style="height:221px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Inferring_LUCA%27s_genome.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Inferring_LUCA%27s_genome.svg/274px-Inferring_LUCA%27s_genome.svg.png" decoding="async" width="274" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Inferring_LUCA%27s_genome.svg/411px-Inferring_LUCA%27s_genome.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Inferring_LUCA%27s_genome.svg/548px-Inferring_LUCA%27s_genome.svg.png 2x" data-file-width="684" data-file-height="554" /></a></span></div><div class="thumbcaption">A direct way to infer LUCA's <a href="/wiki/Genome" title="Genome">genome</a> would be to find genes common to all surviving descendants, but little can be learnt by this approach, as there are only about 30 such genes. They are mostly for <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> proteins, proving that LUCA had the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>. Many other LUCA genes have been lost in later lineages over 4 billion years of evolution.<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-0" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></div></div><div class="tsingle" style="width:562px;max-width:562px"><div class="thumbimage" style="height:221px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Three_ways_to_infer_genes_present_in_LUCA.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e7/Three_ways_to_infer_genes_present_in_LUCA.jpg/560px-Three_ways_to_infer_genes_present_in_LUCA.jpg" decoding="async" width="560" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/e7/Three_ways_to_infer_genes_present_in_LUCA.jpg 1.5x" data-file-width="740" data-file-height="293" /></a></span></div><div class="thumbcaption">Three ways to infer genes present in LUCA: universal presence, presence in both the <a href="/wiki/Bacteria" title="Bacteria">Bacterial</a> and <a href="/wiki/Archaea" title="Archaea">Archaean</a> domains, and presence in two <a href="/wiki/Phylum" title="Phylum">phyla</a> in both domains. The first yields as stated only about 30 genes; the second, some 11,000 with <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">lateral gene transfer</a> (LGT) very likely; the third, 355 genes probably in LUCA, since they were found in at least two phyla in both domains, making LGT an unlikely explanation.<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-1" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></div></div></div></div></div> <p>In 2016, Madeline C. Weiss and colleagues genetically analyzed 6.1 million protein-coding genes and 286,514 protein clusters from sequenced <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotic</a> genomes representing many <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic trees</a>, and identified 355 protein clusters that were probably common to the LUCA. The results of their analysis are highly specific, though debated. They depict LUCA as "<a href="/wiki/Anaerobic_organism" title="Anaerobic organism">anaerobic</a>, <a href="/wiki/Carbon_dioxide" title="Carbon dioxide">CO<sub>2</sub></a>-fixing, <a href="/wiki/Hydrogen" title="Hydrogen">H<sub>2</sub></a>-dependent with a <a href="/wiki/Wood%E2%80%93Ljungdahl_pathway" title="Wood–Ljungdahl pathway">Wood–Ljungdahl pathway</a> (the reductive <a href="/wiki/Acetyl-CoA" title="Acetyl-CoA">acetyl-coenzyme A</a> pathway), <a href="/wiki/Nitrogen" title="Nitrogen">N<sub>2</sub></a>-fixing and <a href="/wiki/Thermophile" title="Thermophile">thermophilic</a>. LUCA's biochemistry was replete with <a href="/wiki/Iron(II)_sulfide" title="Iron(II) sulfide">FeS</a> clusters and <a href="/wiki/Radical_(chemistry)" title="Radical (chemistry)">radical</a> reaction mechanisms."<sup id="cite_ref-Weiss_et_al_20162_11-0" class="reference"><a href="#cite_note-Weiss_et_al_20162-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Cofactor_(biochemistry)" title="Cofactor (biochemistry)">cofactors</a> also reveal "dependence upon <a href="/wiki/Transition_metal" title="Transition metal">transition metals</a>, <a href="/wiki/Flavin_mononucleotide" title="Flavin mononucleotide">flavins</a>, <a href="/wiki/S-adenosyl_methionine" class="mw-redirect" title="S-adenosyl methionine">S-adenosyl methionine</a>, <a href="/wiki/Coenzyme_A" title="Coenzyme A">coenzyme A</a>, <a href="/wiki/Ferredoxin" title="Ferredoxin">ferredoxin</a>, <a href="/wiki/Molybdopterin" title="Molybdopterin">molybdopterin</a>, <a href="/wiki/Corrin" title="Corrin">corrins</a> and <a href="/wiki/Selenium" title="Selenium">selenium</a>. Its genetic code required <a href="/wiki/Nucleoside" title="Nucleoside">nucleoside</a> modifications and S-adenosylmethionine-dependent <a href="/wiki/Methylation" title="Methylation">methylations</a>."<sup id="cite_ref-Weiss_et_al_20162_11-1" class="reference"><a href="#cite_note-Weiss_et_al_20162-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> They show that <a href="/wiki/Methanogen" title="Methanogen">methanogenic</a> <a href="/wiki/Clostridium" title="Clostridium">clostridia</a> were <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal, near the root of the phylogenetic tree</a>, in the 355 protein lineages examined, and that the LUCA may therefore have inhabited an anaerobic <a href="/wiki/Hydrothermal_vent" title="Hydrothermal vent">hydrothermal vent</a> setting in a geochemically active environment rich in H<sub>2</sub>, CO<sub>2</sub>, and iron, where <a href="/wiki/Ocean" title="Ocean">ocean</a> water interacted with hot <a href="/wiki/Magma" title="Magma">magma</a> beneath the <a href="/wiki/Seabed" title="Seabed">ocean floor</a>.<sup id="cite_ref-Weiss_et_al_20162_11-2" class="reference"><a href="#cite_note-Weiss_et_al_20162-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> It is even inferred that LUCA also grew from H<sub>2</sub> and CO<sub>2</sub> via the reverse incomplete Krebs cycle.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Other metabolic pathways inferred in LUCA are the <a href="/wiki/Pentose_phosphate_pathway" title="Pentose phosphate pathway">pentose phosphate pathway</a>, <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a>, and <a href="/wiki/Gluconeogenesis" title="Gluconeogenesis">gluconeogenesis</a>.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Even if phylogenetic evidence may point to a hydrothermal vent environment for a thermophilic LUCA, this does not constitute evidence that the <a href="/wiki/Abiogenesis" title="Abiogenesis">origin of life</a> took place at a hydrothermal vent since mass extinctions may have removed previously existing branches of life.<sup id="cite_ref-Cantine-2017_14-0" class="reference"><a href="#cite_note-Cantine-2017-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>While the gross anatomy of the LUCA can be reconstructed only with much uncertainty, its <a href="/wiki/Metabolic_pathway" title="Metabolic pathway">biochemical mechanisms</a> can be described in some detail, based on the "universal" properties currently shared by all independently living organisms on Earth.<sup id="cite_ref-Wächtershäuser_1998_15-0" class="reference"><a href="#cite_note-Wächtershäuser_1998-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:LUCA_systems_and_environment.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/LUCA_systems_and_environment.svg/440px-LUCA_systems_and_environment.svg.png" decoding="async" width="440" height="319" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/LUCA_systems_and_environment.svg/660px-LUCA_systems_and_environment.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ed/LUCA_systems_and_environment.svg/880px-LUCA_systems_and_environment.svg.png 2x" data-file-width="1008" data-file-height="730" /></a><figcaption>LUCA systems and environment, including the <a href="/wiki/Wood%E2%80%93Ljungdahl_pathway" title="Wood–Ljungdahl pathway">Wood–Ljungdahl or reductive acetyl–CoA pathway</a> to <a href="/wiki/Carbon_fixation" class="mw-redirect" title="Carbon fixation">fix carbon</a>, and most likely <a href="/wiki/DNA" title="DNA">DNA</a> complete with the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> and <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> to <a href="/wiki/DNA_replication" title="DNA replication">replicate</a> it, <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcribe it to RNA</a>, and <a href="/wiki/Translation_(biology)" title="Translation (biology)">translate it to proteins</a>.</figcaption></figure> <p>The LUCA certainly had <a href="/wiki/Gene" title="Gene">genes</a> and a <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>.<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-2" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Its genetic material was most likely DNA,<sup id="cite_ref-Wächtershäuser_1998_15-1" class="reference"><a href="#cite_note-Wächtershäuser_1998-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> so that it lived after the <a href="/wiki/RNA_world" title="RNA world">RNA world</a>.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-PiP_19-0" class="reference"><a href="#cite_note-PiP-19"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The DNA was kept double-stranded by an <a href="/wiki/Enzyme" title="Enzyme">enzyme</a>, <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a>, which recognises the structure and directionality of DNA.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> The integrity of the DNA was maintained by a group of <a href="/wiki/DNA_repair" title="DNA repair">repair</a> enzymes including <a href="/wiki/DNA_topoisomerase" class="mw-redirect" title="DNA topoisomerase">DNA topoisomerase</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> If the genetic code was based on <a href="/wiki/Nucleic_acid_double_helix" title="Nucleic acid double helix">dual-stranded DNA</a>, it was expressed by copying the information to single-stranded RNA. The RNA was produced by a DNA-dependent <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> using nucleotides similar to those of DNA.<sup id="cite_ref-Wächtershäuser_1998_15-2" class="reference"><a href="#cite_note-Wächtershäuser_1998-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> It had multiple <a href="/wiki/DNA-binding_protein" title="DNA-binding protein">DNA-binding proteins</a>, such as histone-fold proteins.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> The genetic code was expressed into <a href="/wiki/Protein" title="Protein">proteins</a>. These were assembled from 20 free <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> by <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> of a <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> via a mechanism of <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>, <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNAs</a>, and a group of related proteins.<sup id="cite_ref-Wächtershäuser_1998_15-3" class="reference"><a href="#cite_note-Wächtershäuser_1998-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>LUCA was likely capable of <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual interaction</a> in the sense that adaptive gene functions were present that promoted the transfer of DNA between individuals of the population to facilitate <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>. Homologous gene products that promote genetic recombination are present in bacteria, archaea and eukaryota, such as the <a href="/wiki/RecA" title="RecA">RecA</a> protein in bacteria, the RadA protein in archaea, and the <a href="/wiki/RAD51" title="RAD51">Rad51</a> and <a href="/wiki/DMC1_(gene)" title="DMC1 (gene)">Dmc1</a> proteins in eukaryota.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The functionality of LUCA as well as evidence for the early evolution membrane-dependent biological systems together suggest that LUCA had cellularity and cell membranes.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> As for the cell's gross structure, it contained a water-based <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> effectively enclosed by a <a href="/wiki/Lipid_bilayer" title="Lipid bilayer">lipid bilayer</a> membrane; it was capable of reproducing by cell division.<sup id="cite_ref-Wächtershäuser_1998_15-4" class="reference"><a href="#cite_note-Wächtershäuser_1998-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> It tended to exclude <a href="/wiki/Sodium" title="Sodium">sodium</a> and concentrate <a href="/wiki/Potassium" title="Potassium">potassium</a> by means of specific <a href="/wiki/Ion_transporter" title="Ion transporter">ion transporters</a> (or ion pumps). The cell multiplied by duplicating all its contents followed by <a href="/wiki/Cellular_division" class="mw-redirect" title="Cellular division">cellular division</a>. The cell used <a href="/wiki/Chemiosmosis" title="Chemiosmosis">chemiosmosis</a> to produce energy. It also <a href="/wiki/Redox" title="Redox">reduced</a> CO<sub>2</sub> and oxidized H<sub>2</sub> (<a href="/wiki/Methanogenesis" title="Methanogenesis">methanogenesis</a> or <a href="/wiki/Acetogenesis" title="Acetogenesis">acetogenesis</a>) via <a href="/wiki/Acetyl" class="mw-redirect" title="Acetyl">acetyl</a>-<a href="/wiki/Thioester" title="Thioester">thioesters</a>.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>By <a href="/wiki/Phylogenetic_bracketing" title="Phylogenetic bracketing">phylogenetic bracketing</a>, analysis of the presumed LUCA's offspring groups, LUCA appears to have been a small, single-celled organism. It likely had a ring-shaped coil of <a href="/wiki/DNA" title="DNA">DNA</a> floating freely within the cell. Morphologically, it would likely not have stood out within a mixed population of small modern-day bacteria. The originator of the <a href="/wiki/Three-domain_system" title="Three-domain system">three-domain system</a>, <a href="/wiki/Carl_Woese" title="Carl Woese">Carl Woese</a>, stated that in its genetic machinery, the LUCA would have been a "simpler, more rudimentary entity than the individual ancestors that spawned the three [domains] (and their descendants)".<sup id="cite_ref-Woese_Kandler_Wheelis_1990_1-1" class="reference"><a href="#cite_note-Woese_Kandler_Wheelis_1990-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Reduktiver_Acetyl-CoA-Weg.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Reduktiver_Acetyl-CoA-Weg.png/350px-Reduktiver_Acetyl-CoA-Weg.png" decoding="async" width="350" height="289" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Reduktiver_Acetyl-CoA-Weg.png/525px-Reduktiver_Acetyl-CoA-Weg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Reduktiver_Acetyl-CoA-Weg.png/700px-Reduktiver_Acetyl-CoA-Weg.png 2x" data-file-width="3648" data-file-height="3007" /></a><figcaption>The LUCA used the <a href="/wiki/Wood%E2%80%93Ljungdahl_pathway" title="Wood–Ljungdahl pathway">Wood–Ljungdahl or reductive acetyl–CoA pathway</a> to <a href="/wiki/Carbon_fixation" class="mw-redirect" title="Carbon fixation">fix carbon</a>, if it was an <a href="/wiki/Autotroph" title="Autotroph">autotroph</a>, or to <a href="/wiki/Anaerobic_respiration" title="Anaerobic respiration">respire anaerobically</a>, if it was a <a href="/wiki/Heterotroph" title="Heterotroph">heterotroph</a>.</figcaption></figure> <p>An alternative to the search for "universal" traits is to use genome analysis to identify phylogenetically ancient genes. This gives a picture of a LUCA that could live in a geochemically harsh environment and is like modern prokaryotes. Analysis of biochemical pathways implies the same sort of chemistry as does phylogenetic analysis. Weiss and colleagues write that "Experiments ... demonstrate that ... <a href="/wiki/Wood%E2%80%93Ljungdahl_pathway" title="Wood–Ljungdahl pathway">acetyl-CoA pathway</a> [chemicals used in anaerobic respiration] <a href="/wiki/Formate" title="Formate">formate</a>, <a href="/wiki/Methanol" title="Methanol">methanol</a>, <a href="/wiki/Acetyl_group" title="Acetyl group">acetyl</a> moieties, and even <a href="/wiki/Pyruvate" class="mw-redirect" title="Pyruvate">pyruvate</a> arise spontaneously ... from CO<sub>2</sub>, native metals, and water", a combination present in hydrothermal vents.<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-3" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>An experiment shows that Zn<sup>2+</sup>, Cr<sup>3+</sup>, and Fe can promote 6 of the 11 reactions of an ancient anabolic pathway called the <a href="/wiki/Reverse_Krebs_cycle" title="Reverse Krebs cycle">reverse Krebs cycle</a> in acidic conditions which implies that LUCA might have inhabited either hydrothermal vents or acidic metal-rich hydrothermal fields.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>Because both bacteria and archaea have differences in the structure of phospholipids and cell wall, ion pumping, most proteins involved in DNA replication, and glycolysis, it is inferred that LUCA had a permeable membrane without an ion pump. The emergence of Na<sup>+</sup>/H<sup>+</sup> antiporters likely lead to the evolution of impermeable membranes present in eukaryotes, archaea, and bacteria. It is stated that "The late and independent evolution of glycolysis but not gluconeogenesis is entirely consistent with LUCA being powered by natural proton gradients across leaky membranes. Several discordant traits are likely to be linked to the late evolution of cell membranes, notably the cell wall, whose synthesis depends on the membrane and DNA replication".<sup id="cite_ref-Sojo_Pomiankowski_Lane_20142_28-0" class="reference"><a href="#cite_note-Sojo_Pomiankowski_Lane_20142-28"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Although LUCA likely had DNA, it is unknown if it could replicate DNA and is suggested to "might just have been a chemically stable repository for RNA-based replication".<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-4" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> It is likely that the permeable membrane of LUCA was composed of archaeal lipids (<a href="/wiki/Isoprenoids" class="mw-redirect" title="Isoprenoids">isoprenoids</a>) and bacterial lipids (<a href="/wiki/Fatty_acid" title="Fatty acid">fatty acids</a>). Isoprenoids would have enhanced stabilization of LUCA's membrane in the surrounding extreme habitat. Nick Lane and coauthors state that "The advantages and disadvantages of incorporating isoprenoids into cell membranes in different microenvironments may have driven membrane divergence, with the later biosynthesis of phospholipids giving rise to the unique G1P and G3P headgroups of archaea and bacteria respectively. If so, the properties conferred by membrane isoprenoids place the lipid divide as early as the <a href="/wiki/Origin_of_life" class="mw-redirect" title="Origin of life">origin of life</a>".<sup id="cite_ref-Jordan_Nee_Lane_20192_29-0" class="reference"><a href="#cite_note-Jordan_Nee_Lane_20192-29"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>A 2024 study suggests that LUCA's genome was similar in size to that of modern prokaryotes, coding for some 2,600 proteins; that it respired anaerobically, and was an <a href="/wiki/Acetogen" title="Acetogen">acetogen</a>; and that it had an early <a href="/wiki/CRISPR" title="CRISPR">CAS</a>-based anti-viral immune system.<sup id="cite_ref-Moody_et_al_2024_30-0" class="reference"><a href="#cite_note-Moody_et_al_2024-30"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alternative_interpretations">Alternative interpretations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=4" title="Edit section: Alternative interpretations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some other researchers have challenged Weiss et al.'s 2016 conclusions. Sarah Berkemer and Shawn McGlynn argue that Weiss et al. undersampled the families of proteins, so that the phylogenetic trees were not complete and failed to describe the evolution of proteins correctly. There are two risks in attempting to attribute LUCA's environment from near-universal gene distribution (as in Weiss et al. 2016). On the one hand, it risks misattributing <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergence</a> or horizontal gene transfer events to vertical descent; on the other hand, it risks misattributing potential LUCA gene families as horizontal gene transfer events. A phylogenomic and geochemical analysis of a set of proteins that probably traced to the LUCA show that it had K<sup>+</sup>-dependent GTPases and the ionic composition and concentration of its intracellular fluid was seemingly high K<sup>+</sup>/Na<sup>+</sup> ratio, <span class="chemf nowrap">NH<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:-0.4em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline">4</sub></span></span></span>, Fe<sup>2+</sup>, CO<sup>2+</sup>, Ni<sup>2+</sup>, Mg<sup>2+</sup>, Mn<sup>2+</sup>, Zn<sup>2+</sup>, pyrophosphate, and <span class="chemf nowrap">PO<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:-0.4em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">3−</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline">4</sub></span></span></span> which would imply a terrestrial <a href="/wiki/Hot_spring" title="Hot spring">hot spring</a> habitat. It possibly had a phosphate-based metabolism. Further, these proteins were unrelated to <a href="/wiki/Autotroph" title="Autotroph">autotrophy</a> (the ability of an organism to create its own <a href="/wiki/Organic_matter" title="Organic matter">organic matter</a>), suggesting that the LUCA had a <a href="/wiki/Heterotroph" title="Heterotroph">heterotrophic</a> lifestyle (consuming organic matter) and that its growth was dependent on organic matter produced by the physical environment.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Nick_Lane" title="Nick Lane">Nick Lane</a> argues that Na<sup>+</sup>/H<sup>+</sup> antiporters could readily explain the low concentration of Na<sup>+</sup> in the LUCA and its descendants. </p><p>The presence of the energy-handling enzymes <a href="/wiki/CODH" class="mw-redirect" title="CODH">CODH</a>/<a href="/wiki/Acetyl-CoA" title="Acetyl-CoA">acetyl-coenzyme A</a> synthase in LUCA could be compatible not only with being an <a href="/wiki/Autotroph" title="Autotroph">autotroph</a> but also with life as a <a href="/wiki/Mixotroph" title="Mixotroph">mixotroph</a> or <a href="/wiki/Heterotroph" title="Heterotroph">heterotroph</a>.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Weiss et al. 2018 reply that no enzyme defines a trophic lifestyle, and that heterotrophs evolved from autotrophs.<sup id="cite_ref-Weiss_Preiner_Xavier_2018_10-5" class="reference"><a href="#cite_note-Weiss_Preiner_Xavier_2018-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Evidence_that_LUCA_was_mesophilic">Evidence that LUCA was mesophilic</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=5" title="Edit section: Evidence that LUCA was mesophilic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Several lines of evidence now suggest that LUCA was non-thermophilic. </p><p>The content of G + C nucleotide pairs (compared to the occurrence of A + T pairs) can indicate an organism's thermal optimum as they are more thermally stable due to an additional hydrogen bond. As a result they occur more frequently in the rRNA of thermophiles; however this is not seen in LUCA's reconstructed rRNA.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cantine-2017_14-1" class="reference"><a href="#cite_note-Cantine-2017-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>The identification of thermophilic genes in the LUCA has been criticized,<sup id="cite_ref-GogartenDeamer20162_35-0" class="reference"><a href="#cite_note-GogartenDeamer20162-35"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> as they may instead represent genes that evolved later in archaea or bacteria, then migrated between these via <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>, as in Woese's 1998 hypothesis.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> For instance, the thermophile-specific topoisomerase, <a href="/wiki/Reverse_gyrase" title="Reverse gyrase">reverse gyrase</a>, was initially attributed to LUCA<sup id="cite_ref-Weiss_et_al_20162_11-3" class="reference"><a href="#cite_note-Weiss_et_al_20162-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> before an exhaustive phylogenetic study revealed a more recent origin of this enzyme followed by extensive horizontal gene transfer.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> LUCA could have been a mesophile that fixed CO<sub>2</sub> and relied on H<sub>2</sub>, and lived close to hydrothermal vents.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>Further evidence that LUCA was <a href="/wiki/Mesophile" title="Mesophile">mesophilic</a> comes from the amino acid composition of its proteins. The abundance of I, V, Y, W, R, E, and L amino acids (denoted IVYWREL) in an organism's proteins is correlated with its optimal growth temperature.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> According to phylogentic analysis, the IVYWREL content of LUCA's proteins suggests its ideal temperature was below 50°C.<sup id="cite_ref-Cantine-2017_14-2" class="reference"><a href="#cite_note-Cantine-2017-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>Finally, evidence that bacteria and archaea both independently underwent phases of increased and subsequently decreased thermo-tolerance suggests a dramatic post-LUCA climate shift that affected both populations and would explain the seeming genetic pervasiveness of thermo-tolerant genetics.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Age">Age</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=6" title="Edit section: Age"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></div> <p>Studies from 2000 to 2018 have suggested an increasingly ancient time for the LUCA. In 2000, estimates of the LUCA's age ranged from 3.5 to 3.8 billion years ago in the <a href="/wiki/Paleoarchean" title="Paleoarchean">Paleoarchean</a>,<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> a few hundred million years before the <a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">earliest fossil evidence of life</a>, for which candidates range in age from 3.48 to 4.28 billion years ago.<sup id="cite_ref-AST-20131108_42-0" class="reference"><a href="#cite_note-AST-20131108-42"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NG-20131208_43-0" class="reference"><a href="#cite_note-NG-20131208-43"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> This placed the origin of the first forms of life shortly after the <a href="/wiki/Late_Heavy_Bombardment" title="Late Heavy Bombardment">Late Heavy Bombardment</a> which was thought to have repeatedly sterilized Earth's surface. However, a 2018 study by Holly Betts and colleagues applied a <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a> model to the genomic and fossil record (102 species, 29 common protein-coding genes, mostly ribosomal), concluding that LUCA preceded the Late Heavy Bombardment (making the LUCA over 3.9 billion years ago).<sup id="cite_ref-Betts_Puttick_Clark_2018_47-0" class="reference"><a href="#cite_note-Betts_Puttick_Clark_2018-47"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> A 2022 study suggested an age of around 3.6-4.2 billion years for the LUCA.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> A 2024 study suggested that the LUCA lived around 4.2 billion years ago (with a confidence interval of 4.09–4.33 billion years ago).<sup id="cite_ref-Moody_et_al_2024_30-1" class="reference"><a href="#cite_note-Moody_et_al_2024-30"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Root_of_the_tree_of_life">Root of the tree of life</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=7" title="Edit section: Root of the tree of life"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Tree_Of_Life_(with_horizontal_gene_transfer).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/220px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/330px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/440px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 2x" data-file-width="1782" data-file-height="2370" /></a><figcaption>2005 <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a> showing <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfers</a> between branches including (coloured lines) the <a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a> of <a href="/wiki/Plastid" title="Plastid">plastids</a> and <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a>. "Horizontal gene transfer and how it has impacted the evolution of life is presented through a web connecting bifurcating branches that complicate, yet do not erase, the tree of life".<sup id="cite_ref-Smets_Barkay_2005_49-0" class="reference"><a href="#cite_note-Smets_Barkay_2005-49"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup></figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For branching of bacteria phyla, see <a href="/wiki/Bacterial_phyla" title="Bacterial phyla">Bacterial phyla</a>.</div> <p>In 1990, a novel concept of the <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a> was presented, dividing the living world into three stems, classified as the domains <a href="/wiki/Bacteria" title="Bacteria">Bacteria</a>, <a href="/wiki/Archaea" title="Archaea">Archaea</a>, <a href="/wiki/Eukarya" class="mw-redirect" title="Eukarya">Eukarya</a>.<sup id="cite_ref-Woese_Kandler_Wheelis_1990_1-2" class="reference"><a href="#cite_note-Woese_Kandler_Wheelis_1990-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Sapp2009_50-0" class="reference"><a href="#cite_note-Sapp2009-50"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Brock_2015_51-0" class="reference"><a href="#cite_note-Brock_2015-51"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Brock_2022_52-0" class="reference"><a href="#cite_note-Brock_2022-52"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> It is the first tree founded exclusively on molecular phylogenetics, and which includes the evolution of microorganisms. It has been called a "universal phylogenetic tree in rooted form".<sup id="cite_ref-Woese_Kandler_Wheelis_1990_1-3" class="reference"><a href="#cite_note-Woese_Kandler_Wheelis_1990-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This tree and its rooting became the subject of debate.<sup id="cite_ref-Sapp2009_50-1" class="reference"><a href="#cite_note-Sapp2009-50"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> </p><p>In the meantime, numerous modifications of this tree, mainly concerning the role and importance of horizontal gene transfer for its rooting and early ramifications have been suggested (e.g.<sup id="cite_ref-Brown_Doolittle_1995_55-0" class="reference"><a href="#cite_note-Brown_Doolittle_1995-55"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smets_Barkay_2005_49-1" class="reference"><a href="#cite_note-Smets_Barkay_2005-49"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup>). Since heredity occurs both vertically and horizontally, the tree of life may have been more weblike or netlike in its early phase and more treelike when it grew three-stemmed.<sup id="cite_ref-Smets_Barkay_2005_49-2" class="reference"><a href="#cite_note-Smets_Barkay_2005-49"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Presumably horizontal gene transfer has decreased with growing cell stability.<sup id="cite_ref-Harold_2014_56-0" class="reference"><a href="#cite_note-Harold_2014-56"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>A modified version of the tree, based on several molecular studies, has its root between a <a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a> <a href="/wiki/Domain_(biology)" title="Domain (biology)">domain</a> <a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> and a <a href="/wiki/Clade" title="Clade">clade</a> formed by <a href="/wiki/Archaea" title="Archaea">Archaea</a> and <a href="/wiki/Eukaryota" class="mw-redirect" title="Eukaryota">Eukaryota</a>.<sup id="cite_ref-Brown_Doolittle_1995_55-1" class="reference"><a href="#cite_note-Brown_Doolittle_1995-55"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> A small minority of studies place the root in the domain bacteria, in the phylum <a href="/wiki/Bacillota" title="Bacillota">Bacillota</a>,<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> or state that the phylum <a href="/wiki/Chloroflexota" title="Chloroflexota">Chloroflexota</a> (formerly Chloroflexi) is <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal</a> to a clade with Archaea and Eukaryotes and the rest of bacteria (as proposed by <a href="/wiki/Thomas_Cavalier-Smith" title="Thomas Cavalier-Smith">Thomas Cavalier-Smith</a>).<sup id="cite_ref-CS2_58-0" class="reference"><a href="#cite_note-CS2-58"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Metagenomics" title="Metagenomics">Metagenomic</a> analyses recover a two-domain system with the domains Archaea and Bacteria; in this view of the tree of life, Eukaryotes are derived from Archaea.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> With the later gene pool of LUCA's descendants, sharing a common framework of the <a href="/wiki/Chargaff%27s_rules" title="Chargaff's rules">AT/GC rule</a> and the standard twenty amino acids, horizontal gene transfer would have become feasible and could have been common.<sup id="cite_ref-Harris_Hill_2021_62-0" class="reference"><a href="#cite_note-Harris_Hill_2021-62"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p><p>The nature of LUCA remains disputed. In 1994, on the basis of primordial metabolism (sensu <a href="/wiki/G%C3%BCnter_W%C3%A4chtersh%C3%A4user" title="Günter Wächtershäuser">Wächtershäuser</a>), <a href="/wiki/Otto_Kandler" title="Otto Kandler">Otto Kandler</a> proposed a successive divergence of the three domains of life<sup id="cite_ref-Woese_Kandler_Wheelis_1990_1-4" class="reference"><a href="#cite_note-Woese_Kandler_Wheelis_1990-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> from a multiphenotypical <i>population</i> of <a href="/wiki/Pre-cell" title="Pre-cell">pre-cells</a>, reached by gradual evolutionary improvements (<a href="/wiki/Cellularization" title="Cellularization">cellularization</a>).<sup id="cite_ref-Kandler_1994_63-0" class="reference"><a href="#cite_note-Kandler_1994-63"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kandler_1995_64-0" class="reference"><a href="#cite_note-Kandler_1995-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kandler_1998_65-0" class="reference"><a href="#cite_note-Kandler_1998-65"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> These phenotypically diverse pre-cells were metabolising, self-reproducing entities exhibiting frequent mutual exchange of genetic information. Thus, in this scenario there was no "first cell". It may explain the unity and, at the same time, the partition into three lines (the three domains) of life. Kandler's pre-cell theory is supported by Wächtershäuser.<sup id="cite_ref-Wächtershäuser_2003_66-0" class="reference"><a href="#cite_note-Wächtershäuser_2003-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wächtershäuser_2006_67-0" class="reference"><a href="#cite_note-Wächtershäuser_2006-67"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> In 1998, <a href="/wiki/Carl_Woese" title="Carl Woese">Carl Woese</a>, based on the RNA world concept, proposed that no individual organism could be considered a LUCA, and that the genetic heritage of all modern organisms derived through <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a> among an ancient community of organisms.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Other authors concur that there was a "complex collective genome"<sup id="cite_ref-Egel_2012_69-0" class="reference"><a href="#cite_note-Egel_2012-69"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> at the time of the LUCA, and that horizontal gene transfer was important in the evolution of later groups;<sup id="cite_ref-Egel_2012_69-1" class="reference"><a href="#cite_note-Egel_2012-69"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> Nicolas Glansdorff states that LUCA "was in a metabolically and morphologically heterogeneous community, constantly shuffling around genetic material" and "remained an evolutionary entity, though loosely defined and constantly changing, as long as this promiscuity lasted."<sup id="cite_ref-Glansdorff_Xu_Labedan_2008_70-0" class="reference"><a href="#cite_note-Glansdorff_Xu_Labedan_2008-70"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p><p>The theory of a universal common ancestry of life is widely accepted. In 2010, based on "the vast array of molecular sequences now available from all domains of life,"<sup id="cite_ref-Steel_71-0" class="reference"><a href="#cite_note-Steel-71"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> D. L. Theobald published a "<a href="/wiki/Statistical_hypothesis_test" title="Statistical hypothesis test">formal test</a>" of universal common ancestry (UCA). This deals with the <a href="/wiki/Common_descent" title="Common descent">common descent</a> of all extant terrestrial organisms, each being a genealogical descendant of a single species from the distant past. His formal test favoured the existence of a universal common ancestry over a wide class of alternative hypotheses that included horizontal gene transfer. Basic biochemical principles imply that all organisms do have a common ancestry.<sup id="cite_ref-Theobald-2010_72-0" class="reference"><a href="#cite_note-Theobald-2010-72"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p><p>A proposed, earlier, non-cellular ancestor to LUCA is the <a href="/wiki/First_universal_common_ancestor" title="First universal common ancestor">First universal common ancestor</a> (FUCA).<sup id="cite_ref-Prosdocimi_2019_73-0" class="reference"><a href="#cite_note-Prosdocimi_2019-73"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Prosdocimi_2023_74-0" class="reference"><a href="#cite_note-Prosdocimi_2023-74"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> FUCA would therefore be the ancestor to every modern cell as well as ancient, now-extinct cellular lineages not descendant of LUCA. FUCA is assumed to have had other descendants than LUCA, none of which have modern descendants. Some genes of these ancient now-extinct cell lineages are thought to have been <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontally transferred</a> into the genome of early descendants of LUCA.<sup id="cite_ref-Harris_Hill_2021_62-1" class="reference"><a href="#cite_note-Harris_Hill_2021-62"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="LUCA_and_viruses">LUCA and viruses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=8" title="Edit section: LUCA and viruses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Viral_evolution" title="Viral evolution">origin of viruses</a> remains disputed. Since viruses need host cells for their replication, it is likely that they emerged <i>after</i> the <a href="/wiki/Evolution_of_cells" title="Evolution of cells">formation of cells</a>. Viruses may even have multiple origins and different types of viruses may have evolved independently over the history of life.<sup id="cite_ref-Brock_2022_52-1" class="reference"><a href="#cite_note-Brock_2022-52"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> There are different hypotheses for the origins of viruses, for instance an early viral origin from the <a href="/wiki/RNA_world" title="RNA world">RNA world</a> or a later viral origin from <a href="/wiki/Selfish_dna" class="mw-redirect" title="Selfish dna">selfish DNA</a>.<sup id="cite_ref-Brock_2022_52-2" class="reference"><a href="#cite_note-Brock_2022-52"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>Based on how viruses are currently distributed across the <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>, the LUCA is suspected of having been prey to multiple viruses, ancestral to those that now have those two domains as their hosts.<sup id="cite_ref-KrupovicLUCA2020_75-0" class="reference"><a href="#cite_note-KrupovicLUCA2020-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Furthermore, extensive virus evolution seems to have preceded the LUCA, since the <a href="/wiki/Jelly_roll_fold" title="Jelly roll fold">jelly-roll structure</a> of <a href="/wiki/Capsid" title="Capsid">capsid</a> proteins is shared by RNA and DNA viruses across all three domains of life.<sup id="cite_ref-Forterre_Prangishvili_2009_76-0" class="reference"><a href="#cite_note-Forterre_Prangishvili_2009-76"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Durzyńska_Goździcka-Józefiak_2015_77-0" class="reference"><a href="#cite_note-Durzyńska_Goździcka-Józefiak_2015-77"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> LUCA's viruses were probably mainly dsDNA viruses in the groups called <i><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i> and <i><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></i>. Two other <a href="/wiki/Single-stranded_DNA_virus" class="mw-redirect" title="Single-stranded DNA virus">single-stranded DNA virus</a> groups within the <i><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></i>, the <i><a href="/wiki/Microviridae" title="Microviridae">Microviridae</a></i> and the <i><a href="/wiki/Tubulavirales" title="Tubulavirales">Tubulavirales</a></i>, likely infected the last bacterial common ancestor. The last archaeal common ancestor was probably host to spindle-shaped viruses. All of these could well have affected the LUCA, in which case each must since have been lost in the host domain where it is no longer extant. By contrast, RNA viruses do not appear to have been important parasites of LUCA, even though straightforward thinking might have envisaged viruses as beginning with <a href="/wiki/RNA_viruses" class="mw-redirect" title="RNA viruses">RNA viruses</a> directly derived from an RNA world. Instead, by the time the LUCA lived, RNA viruses had probably already been out-competed by DNA viruses.<sup id="cite_ref-KrupovicLUCA2020_75-1" class="reference"><a href="#cite_note-KrupovicLUCA2020-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> </p><p>LUCA might have been the ancestor to some viruses, as it might have had at least two descendants: LUCELLA, the Last Universal Cellular Ancestor, the ancestor to all cells, and the archaic virocell ancestor, the ancestor to large-to-medium-sized <a href="/wiki/DNA_virus" title="DNA virus">DNA viruses</a>.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> Viruses might have evolved before LUCA but after the <a href="/wiki/First_universal_common_ancestor" title="First universal common ancestor">First universal common ancestor</a> (FUCA), according to the reduction hypothesis, where <a href="/wiki/Giant_virus" title="Giant virus">giant viruses</a> evolved from primordial cells that became <a href="/wiki/Parasitism" title="Parasitism">parasitic</a>.<sup id="cite_ref-Harris_Hill_2021_62-2" class="reference"><a href="#cite_note-Harris_Hill_2021-62"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=9" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a> – Life arising from non-living matter</li> <li><a href="/wiki/Cellularization" title="Cellularization">Cellularization</a> – Scientific theory to explain the origin and formation of cells</li> <li><a href="/wiki/Chemoton" title="Chemoton">Chemoton</a> – Abstract model for the fundamental unit of life</li> <li><a href="/wiki/Darwinian_threshold" title="Darwinian threshold">Darwinian threshold</a> – Period during the evolution of the first cells</li> <li><a href="/wiki/Last_eukaryotic_common_ancestor" class="mw-redirect" title="Last eukaryotic common ancestor">Last eukaryotic common ancestor</a> – Process of forming the first eukaryotic cell<span style="display:none" class="category-annotation-with-redirected-description">Pages displaying short descriptions of redirect targets</span></li> <li><a href="/wiki/Mitochondrial_Eve" title="Mitochondrial Eve">Mitochondrial Eve</a> – Matrilineal most recent common ancestor of all living humans</li> <li><a href="/wiki/Pre-cell" title="Pre-cell">Pre-cell</a> – Hypothetical life before complete cells</li> <li><a href="/wiki/Proto-metabolism" title="Proto-metabolism">Proto-metabolism</a> – Chemical reactions which turn into modern metabolism</li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of the evolutionary history of life</a></li> <li><a href="/wiki/Urmetazoan" title="Urmetazoan">Urmetazoan</a> – Hypothetical last common ancestor of all animals</li> <li><a href="/wiki/Y-chromosomal_Adam" title="Y-chromosomal Adam">Y-chromosomal Adam</a> – Patrilineal most recent common ancestor of all living humans</li></ul> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=10" title="Edit section: Notes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text">Other studies propose that LUCA may have been defined wholly through <a href="/wiki/RNA" title="RNA">RNA</a>,<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> consisted of a RNA-DNA hybrid genome, or possessed a retrovirus-like genetic cycle with DNA serving as a stable genetic repository.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-54"><span class="mw-cite-backlink"><b><a href="#cite_ref-54">^</a></b></span> <span class="reference-text">One debate dealt with a former <a href="/wiki/Cladistics" title="Cladistics">cladistic</a> hypothesis: The tree could not be ascribed a root in the usual algorithmic way, because that would require an <a href="/wiki/Outgroup_(cladistics)" title="Outgroup (cladistics)">outgroup</a> for reference. In the case of the universal tree, no outgroup would exist. The cladistic method was used "to root the purple bacteria, for example. But establishing a root for the universal tree of life, the branching order among the primary urkingdoms, was another matter entirely."<sup id="cite_ref-FOOTNOTESapp2009255_53-0" class="reference"><a href="#cite_note-FOOTNOTESapp2009255-53"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Last_universal_common_ancestor&action=edit&section=11" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist reflist-columns references-column-width" style="column-width: 25em;"> <ol class="references"> <li id="cite_note-Woese_Kandler_Wheelis_1990-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Woese_Kandler_Wheelis_1990_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Woese_Kandler_Wheelis_1990_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Woese_Kandler_Wheelis_1990_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Woese_Kandler_Wheelis_1990_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Woese_Kandler_Wheelis_1990_1-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFWoeseKandlerWheelis1990" class="citation journal cs1"><a href="/wiki/Carl_Woese" title="Carl Woese">Woese, C.R.</a>; 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J.; Pisani, Davide (2018). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6152910">"Integrated genomic and fossil evidence illuminates life's early evolution and eukaryote origin"</a>. <i>Nature Ecology & Evolution</i>. <b>2</b> (10): 1556–1562. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018NatEE...2.1556B">2018NatEE...2.1556B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41559-018-0644-x">10.1038/s41559-018-0644-x</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6152910">6152910</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/30127539">30127539</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Ecology+%26+Evolution&rft.atitle=Integrated+genomic+and+fossil+evidence+illuminates+life%27s+early+evolution+and+eukaryote+origin&rft.volume=2&rft.issue=10&rft.pages=1556-1562&rft.date=2018&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6152910%23id-name%3DPMC&rft_id=info%3Apmid%2F30127539&rft_id=info%3Adoi%2F10.1038%2Fs41559-018-0644-x&rft_id=info%3Abibcode%2F2018NatEE...2.1556B&rft.aulast=Betts&rft.aufirst=Holly+C.&rft.au=Puttick%2C+Mark+N.&rft.au=Clark%2C+James+W.&rft.au=Williams%2C+Tom+A.&rft.au=Donoghue%2C+Philip+C.+J.&rft.au=Pisani%2C+Davide&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6152910&rfr_id=info%3Asid%2Fen.wikipedia.org%3ALast+universal+common+ancestor" class="Z3988"></span></span> </li> <li id="cite_note-48"><span class="mw-cite-backlink"><b><a href="#cite_ref-48">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoodyMahendrarajahDombrowskiClark2022" class="citation journal cs1">Moody, Edmund RR; Mahendrarajah, Tara A.; Dombrowski, Nina; Clark, James W.; Petitjean, Celine; Offre, Pierre; Szöllősi, Gergely J.; Spang, Anja; Williams, Tom A. (22 February 2022). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8890751">"An estimate of the deepest branches of the tree of life from ancient vertically evolving genes"</a>. <i>eLife</i>. <b>11</b>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.7554%2FeLife.66695">10.7554/eLife.66695</a></span>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2050-084X">2050-084X</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8890751">8890751</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/35190025">35190025</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=eLife&rft.atitle=An+estimate+of+the+deepest+branches+of+the+tree+of+life+from+ancient+vertically+evolving+genes&rft.volume=11&rft.date=2022-02-22&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8890751%23id-name%3DPMC&rft.issn=2050-084X&rft_id=info%3Apmid%2F35190025&rft_id=info%3Adoi%2F10.7554%2FeLife.66695&rft.aulast=Moody&rft.aufirst=Edmund+RR&rft.au=Mahendrarajah%2C+Tara+A.&rft.au=Dombrowski%2C+Nina&rft.au=Clark%2C+James+W.&rft.au=Petitjean%2C+Celine&rft.au=Offre%2C+Pierre&rft.au=Sz%C3%B6ll%C5%91si%2C+Gergely+J.&rft.au=Spang%2C+Anja&rft.au=Williams%2C+Tom+A.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8890751&rfr_id=info%3Asid%2Fen.wikipedia.org%3ALast+universal+common+ancestor" class="Z3988"></span></span> </li> <li id="cite_note-Smets_Barkay_2005-49"><span class="mw-cite-backlink">^ <a href="#cite_ref-Smets_Barkay_2005_49-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Smets_Barkay_2005_49-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Smets_Barkay_2005_49-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSmetsBarkay2005" class="citation journal cs1">Smets, Barth F.; Barkay, Tamar (September 2005). <a rel="nofollow" class="external text" href="https://www.researchgate.net/publication/7615190">"Horizontal gene transfer: perspectives at a crossroads of scientific disciplines"</a>. <i>Nature Reviews Microbiology</i>. <b>3</b> (9): 675–678. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrmicro1253">10.1038/nrmicro1253</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16145755">16145755</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:2265315">2265315</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews+Microbiology&rft.atitle=Horizontal+gene+transfer%3A+perspectives+at+a+crossroads+of+scientific+disciplines&rft.volume=3&rft.issue=9&rft.pages=675-678&rft.date=2005-09&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2265315%23id-name%3DS2CID&rft_id=info%3Apmid%2F16145755&rft_id=info%3Adoi%2F10.1038%2Fnrmicro1253&rft.aulast=Smets&rft.aufirst=Barth+F.&rft.au=Barkay%2C+Tamar&rft_id=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F7615190&rfr_id=info%3Asid%2Fen.wikipedia.org%3ALast+universal+common+ancestor" class="Z3988"></span></span> </li> <li id="cite_note-Sapp2009-50"><span class="mw-cite-backlink">^ <a href="#cite_ref-Sapp2009_50-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Sapp2009_50-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSapp2009" class="citation book cs1"><a href="/wiki/Jan_Sapp" title="Jan Sapp">Sapp, Jan A.</a> (2009). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=d7zOviXnbSYC"><i>The new foundations of evolution: on the tree of life</i></a>. New York: Oxford University Press. pp. Chapter 19: 257ff (novel concept of the tree of life), Chapters 17-21 plus concluding remarks: 226-318 (discussion of the tree and its rooting), 286ff (LUCA). <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-199-73438-2" title="Special:BookSources/978-0-199-73438-2"><bdi>978-0-199-73438-2</bdi></a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20231106185704/https://books.google.com/books?id=d7zOviXnbSYC">Archived</a> from the original on 6 November 2023<span class="reference-accessdate">. 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Goździcka-Józefiak, Anna (16 October 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4609113">"Viruses and cells intertwined since the dawn of evolution"</a>. <i>Virology Journal</i>. <b>12</b> (1): 169. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1186%2Fs12985-015-0400-7">10.1186/s12985-015-0400-7</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4609113">4609113</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26475454">26475454</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Virology+Journal&rft.atitle=Viruses+and+cells+intertwined+since+the+dawn+of+evolution&rft.volume=12&rft.issue=1&rft.pages=169&rft.date=2015-10-16&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4609113%23id-name%3DPMC&rft_id=info%3Apmid%2F26475454&rft_id=info%3Adoi%2F10.1186%2Fs12985-015-0400-7&rft.aulast=Durzy%C5%84ska&rft.aufirst=Julia&rft.au=Go%C5%BAdzicka-J%C3%B3zefiak%2C+Anna&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4609113&rfr_id=info%3Asid%2Fen.wikipedia.org%3ALast+universal+common+ancestor" class="Z3988"></span></span> </li> <li id="cite_note-78"><span class="mw-cite-backlink"><b><a href="#cite_ref-78">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNasirKimCaetano-Anollés2012" class="citation journal cs1">Nasir, Arshan; 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London: <a href="/wiki/Profile_Books" title="Profile Books">Profile Books</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1781250372" title="Special:BookSources/978-1781250372"><bdi>978-1781250372</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Vital+Question&rft.place=London&rft.pub=Profile+Books&rft.date=2016&rft.isbn=978-1781250372&rft.aulast=Lane&rft.aufirst=Nick&rfr_id=info%3Asid%2Fen.wikipedia.org%3ALast+universal+common+ancestor" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline 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.navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Origin_of_life" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align:center;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar 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href="/wiki/Template:Origin_of_life" title="Template:Origin of life"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Origin_of_life" title="Template talk:Origin of life"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Origin_of_life" title="Special:EditPage/Template:Origin of life"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Origin_of_life" style="font-size:114%;margin:0 4em"><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/History_of_research_into_the_origin_of_life" title="History of research into the origin of life">History of research</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a> (5th century BC)</li> <li><a href="/wiki/Spontaneous_generation" title="Spontaneous generation">Spontaneous generation</a> (4th century BC)</li> <li><a href="/wiki/Primordial_soup" title="Primordial soup">Primordial soup</a> (19th century)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Prebiotic synthesis</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pseudo-panspermia" title="Pseudo-panspermia">Pseudo-panspermia</a></li> <li><a href="/wiki/Miller%E2%80%93Urey_experiment" title="Miller–Urey experiment">Miller–Urey experiment</a></li> <li><a href="/wiki/Formamide-based_prebiotic_chemistry" title="Formamide-based prebiotic chemistry">Formamide-based prebiotic chemistry</a></li> <li><a href="/wiki/Alternative_abiogenesis_scenarios" title="Alternative abiogenesis scenarios">Alternative abiogenesis scenarios</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/Protocell" title="Protocell">Protocells</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/GADV-protein_world_hypothesis" title="GADV-protein world hypothesis">GADV-protein world</a></li> <li><a href="/wiki/Graham_Cairns-Smith#Clay_hypothesis" title="Graham Cairns-Smith">Clay hypothesis</a></li> <li><a href="/wiki/Iron%E2%80%93sulfur_world_hypothesis" title="Iron–sulfur world hypothesis">Iron–sulfur world</a> <ul><li><a href="/wiki/Primordial_sandwich" title="Primordial sandwich">Primordial sandwich</a></li></ul></li> <li><a href="/wiki/PAH_world" class="mw-redirect" title="PAH world">PAH world</a></li> <li><a href="/wiki/Peptide-RNA_world" class="mw-redirect" title="Peptide-RNA world">Peptide-RNA world</a></li> <li><a href="/wiki/Quasispecies_model" title="Quasispecies model">Quasispecies model</a></li> <li><a href="/wiki/RNA_world" title="RNA world">RNA world</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Earliest organisms</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a class="mw-selflink selflink">Last universal common ancestor</a> (LUCA)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Research</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Paleobiology" title="Paleobiology">Paleobiology</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a class="mw-selflink selflink">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Self-replicating_organic_structures" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align: center;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Self-replicating_organic_structures" title="Template:Self-replicating organic structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Self-replicating_organic_structures" title="Template talk:Self-replicating organic structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Self-replicating_organic_structures" title="Special:EditPage/Template:Self-replicating organic structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Self-replicating_organic_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br />agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus<br />dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br />self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br />elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Life,_non-cellular_life,_and_comparable_structures" title="Template:Life, non-cellular life, and comparable structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life,_non-cellular_life,_and_comparable_structures" title="Template talk:Life, non-cellular life, and comparable structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life,_non-cellular_life,_and_comparable_structures" title="Special:EditPage/Template:Life, non-cellular life, and comparable structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Life,_non-cellular_life,_and_comparable_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Life" title="Life">Life</a>, <a href="/wiki/Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">"<a href="/wiki/Prokaryote" title="Prokaryote">Prokaryota</a>"</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li></ul></li> <li><a href="/wiki/LUCA" class="mw-redirect" title="LUCA">LUCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li>'<a href="/wiki/Protist" title="Protist">Protista</a>'</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Virus" title="Virus">Virus</a> <hr /><a href="/wiki/Viroid" title="Viroid">Viroid</a> <hr /><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></i></li> <li><i><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i></li> <li><i><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></i></li> <li><i><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></i></li> <li><i><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></i></li> <li><i><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><i><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></i></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></i></li> <li><i><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></i></li> <li><i><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li> <li><i><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></i></li> <li><i><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></i></li> <li><i><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></i></li> <li><i><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></i></li> <li><i><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></i></li> <li><i><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></i></li> <li><i><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></i></li> <li><i><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></i></li> <li><i><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></i></li> <li><i><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></i></li> <li><i><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></i></li> <li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></i></li> <li><i><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></i></li> <li><i><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></i></li> <li><i><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nanobe" title="Nanobe">Nanobe</a> (?)</li> <li><a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable<br />structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li> <li>"<i><a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></i>"</li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Prion" title="Prion">Prion</a> <ul><li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a> <a href="/wiki/Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li> <li>Retroelements not elsewhere classified <ul><li><a href="/wiki/Retron" title="Retron">Retron</a></li> <li><a href="/wiki/Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li> <li><a href="/wiki/Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li> <li><a href="/wiki/Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Spiegelman%27s_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li> <li><a href="/wiki/Tandem_repeat" 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118.663 11 Template:Navbox"," 8.55% 105.743 2 Template:Efn"," 8.12% 100.492 1 Template:Sfn"," 6.56% 81.101 1 Template:Origin_of_life"," 5.57% 68.935 11 Template:Cite_book"," 4.81% 59.546 1 Template:Short_description"]},"scribunto":{"limitreport-timeusage":{"value":"0.783","limit":"10.000"},"limitreport-memusage":{"value":20886826,"limit":52428800},"limitreport-logs":"anchor_id_list = table#1 {\n [\"CITEREFAdamBorrelGribaldo2018\"] = 1,\n [\"CITEREFAhmadXuWang2017\"] = 1,\n [\"CITEREFBellBoehnkeHarrisonMao2015\"] = 1,\n [\"CITEREFBettsPuttickClarkWilliams2018\"] = 1,\n [\"CITEREFBoussauBlanquartNecsuleaLartillot2008\"] = 1,\n [\"CITEREFBrownDoolittle1995\"] = 1,\n [\"CITEREFCamprubíde_LeeuwHouseRaulin2019\"] = 1,\n [\"CITEREFCantineFournier2017\"] = 1,\n [\"CITEREFCatchpoleForterre2019\"] = 1,\n [\"CITEREFCavalier-Smith2006\"] = 1,\n [\"CITEREFDarwin1859\"] = 1,\n [\"CITEREFDoddPapineauGrenneSlack,_John_F.2017\"] = 1,\n [\"CITEREFDoolittle2000\"] = 1,\n [\"CITEREFDurzyńskaGoździcka-Józefiak2015\"] = 1,\n [\"CITEREFEgel2012\"] = 1,\n [\"CITEREFForterre1997\"] = 1,\n [\"CITEREFForterrePrangishvili2009\"] = 1,\n [\"CITEREFGaltierTourasseGouy1999\"] = 1,\n [\"CITEREFGarwood2012\"] = 1,\n [\"CITEREFGlansdorffXuLabedan2008\"] = 1,\n [\"CITEREFGogartenDeamer2016\"] = 1,\n [\"CITEREFGogartenTaiz1992\"] = 1,\n [\"CITEREFGregory2008\"] = 1,\n [\"CITEREFGroussinBoussauCharlesBlanquart2013\"] = 1,\n [\"CITEREFHarold2014\"] = 1,\n [\"CITEREFHarrisHill2021\"] = 1,\n [\"CITEREFHarrisonLane2018\"] = 1,\n [\"CITEREFHarrisonPalmeiraHalpernLane2022\"] = 1,\n [\"CITEREFHassenkamAnderssonDalbyMackenzie2017\"] = 1,\n [\"CITEREFHugBakerAnantharamanBrown2016\"] = 1,\n [\"CITEREFJordanNeeLane2019\"] = 1,\n [\"CITEREFKandler1994\"] = 1,\n [\"CITEREFKandler1995\"] = 1,\n [\"CITEREFKandler1998\"] = 1,\n [\"CITEREFKooninGalperin2003\"] = 1,\n [\"CITEREFKooninKrupovicIshinoIshino2020\"] = 1,\n [\"CITEREFKooninMartin2005\"] = 1,\n [\"CITEREFKrupovicDoljaKoonin2020\"] = 1,\n [\"CITEREFLamarck1994\"] = 1,\n [\"CITEREFLaneAllenMartin2010\"] = 1,\n [\"CITEREFLupasAlva2018\"] = 1,\n [\"CITEREFMadiganAiyerBuckleySattley2022\"] = 1,\n [\"CITEREFMadiganMartinkoBenderBuckley2015\"] = 1,\n [\"CITEREFMarshall\"] = 1,\n [\"CITEREFMartinRussell2007\"] = 1,\n [\"CITEREFMoodyMahendrarajahDombrowskiClark2022\"] = 1,\n [\"CITEREFMoodyÁlvarez-CarreteroMahendrarajah2024\"] = 1,\n [\"CITEREFMoodyÁlvarez-CarreteroMahendrarajahClark2024\"] = 1,\n [\"CITEREFMuchowskaVarmaChevallot-BerouxLethuillier-Karl2017\"] = 1,\n [\"CITEREFMulkidjanianBychkovDibrovaGalperin2012\"] = 1,\n [\"CITEREFNasirKimCaetano-Anollés2012\"] = 1,\n [\"CITEREFNoble2020\"] = 1,\n [\"CITEREFNoffkeChristianWaceyHazen2013\"] = 1,\n [\"CITEREFOhtomoKakegawaIshidaNagase2013\"] = 1,\n [\"CITEREFProsdocimiJoséde_Farias2019\"] = 2,\n [\"CITEREFRaymannBrochier-ArmanetGribaldo2015\"] = 1,\n [\"CITEREFSapp2009\"] = 1,\n [\"CITEREFSmetsBarkay2005\"] = 1,\n [\"CITEREFSojoPomiankowskiLane2014\"] = 1,\n [\"CITEREFSteelPenny2010\"] = 1,\n [\"CITEREFTheobald2010\"] = 1,\n [\"CITEREFValasBourne2011\"] = 1,\n [\"CITEREFWeissPreinerXavierZimorski2018\"] = 1,\n [\"CITEREFWeissSousaMrnjavacNeukirchen2016\"] = 1,\n [\"CITEREFWikham1995\"] = 1,\n [\"CITEREFWilliamsFosterCoxEmbley2013\"] = 1,\n [\"CITEREFWoese1998\"] = 2,\n [\"CITEREFWoeseKandlerWheelis1990\"] = 1,\n [\"CITEREFWächtershäuser1998\"] = 1,\n [\"CITEREFWächtershäuser2003\"] = 1,\n [\"CITEREFWächtershäuser2006\"] = 1,\n [\"CITEREFZeldovichBerezovskyShakhnovich2007\"] = 1,\n}\ntemplate_list = table#1 {\n [\"!\"] = 1,\n [\"Annotated link\"] = 11,\n [\"Blockquote\"] = 1,\n [\"Chem\"] = 2,\n [\"Circa\"] = 1,\n [\"Citation\"] = 1,\n [\"Cite book\"] = 11,\n [\"Cite journal\"] = 61,\n [\"Cite magazine\"] = 1,\n [\"Cite thesis\"] = 1,\n [\"Creative Commons text attribution notice\"] = 1,\n [\"Efn\"] = 2,\n [\"Evolution\"] = 1,\n [\"For\"] = 1,\n [\"Further\"] = 3,\n [\"Good article\"] = 1,\n [\"Multiple image\"] = 1,\n [\"Notelist\"] = 1,\n [\"Organisms et al.\"] = 1,\n [\"Origin of life\"] = 1,\n [\"ProQuest\"] = 1,\n [\"Redirect\"] = 1,\n [\"Reflist\"] = 1,\n [\"Self-replicating organic structures\"] = 1,\n [\"Sfn\"] = 1,\n [\"Short description\"] = 1,\n [\"Use dmy dates\"] = 1,\n [\"Webarchive\"] = 1,\n}\narticle_whitelist = table#1 {\n}\nciteref_patterns = table#1 {\n}\n"},"cachereport":{"origin":"mw-web.codfw.main-858ddf96d-lxrjw","timestamp":"20241211173550","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Last universal common ancestor","url":"https:\/\/en.wikipedia.org\/wiki\/Last_universal_common_ancestor","sameAs":"http:\/\/www.wikidata.org\/entity\/Q849168","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q849168","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2004-12-20T09:07:06Z","dateModified":"2024-12-06T20:18:27Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/e\/ef\/Phylogenetic_tree_of_life_1990_LUCA.svg","headline":"most recent common ancestor of all current life on Earth"}</script> </body> </html>