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aria-controls="toc-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 Features subsection</span> </button> <ul id="toc-Features-sublist" class="vector-toc-list"> <li id="toc-Reading_frame" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reading_frame"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Reading frame</span> </div> </a> <ul id="toc-Reading_frame-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Start_and_stop_codons" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Start_and_stop_codons"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Start and stop codons</span> </div> </a> <ul id="toc-Start_and_stop_codons-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Effect_of_mutations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Effect_of_mutations"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Effect of mutations</span> </div> </a> <ul id="toc-Effect_of_mutations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Degeneracy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Degeneracy"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Degeneracy</span> </div> </a> <ul id="toc-Degeneracy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Codon_usage_bias" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Codon_usage_bias"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Codon usage bias</span> </div> </a> <ul id="toc-Codon_usage_bias-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Alternative_genetic_codes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Alternative_genetic_codes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Alternative genetic codes</span> </div> </a> <button aria-controls="toc-Alternative_genetic_codes-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 Alternative genetic codes subsection</span> </button> <ul id="toc-Alternative_genetic_codes-sublist" class="vector-toc-list"> <li id="toc-Non-standard_amino_acids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-standard_amino_acids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Non-standard amino acids</span> </div> </a> <ul id="toc-Non-standard_amino_acids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Variations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Variations"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Variations</span> </div> </a> <ul id="toc-Variations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inference" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inference"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Inference</span> </div> </a> <ul id="toc-Inference-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Origin" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Origin"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Origin</span> </div> </a> <ul id="toc-Origin-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">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-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">6</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">7</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>External links</span> </div> </a> <ul id="toc-External_links-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" title="Table of Contents" > <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">Genetic code</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 64 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-64" 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">64 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B4%D9%8A%D9%81%D8%B1%D8%A9_%D8%AC%D9%8A%D9%86%D9%8A%D8%A9" title="شيفرة جينية – Arabic" lang="ar" hreflang="ar" data-title="شيفرة جينية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Genetik_kod" title="Genetik kod – Azerbaijani" lang="az" hreflang="az" data-title="Genetik kod" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AC%E0%A6%82%E0%A6%B6%E0%A6%97%E0%A6%A4%E0%A7%80%E0%A6%AF%E0%A6%BC_%E0%A6%B8%E0%A6%82%E0%A6%95%E0%A7%87%E0%A6%A4" title="বংশগতীয় সংকেত – Bangla" lang="bn" hreflang="bn" data-title="বংশগতীয় সংকেত" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D1%8B%D1%87%D0%BD%D1%8B_%D0%BA%D0%BE%D0%B4" title="Генетычны код – Belarusian" lang="be" hreflang="be" data-title="Генетычны код" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D1%87%D0%B5%D0%BD_%D0%BA%D0%BE%D0%B4" title="Генетичен код – Bulgarian" lang="bg" hreflang="bg" data-title="Генетичен код" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Geneti%C4%8Dki_kod" title="Genetički kod – Bosnian" lang="bs" hreflang="bs" data-title="Genetički kod" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Codi_gen%C3%A8tic" title="Codi genètic – Catalan" lang="ca" hreflang="ca" data-title="Codi genètic" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Genetick%C3%BD_k%C3%B3d" title="Genetický kód – Czech" lang="cs" hreflang="cs" data-title="Genetický kód" 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/Genetisk_kode" title="Genetisk kode – Danish" lang="da" hreflang="da" data-title="Genetisk kode" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-se mw-list-item"><a href="https://se.wikipedia.org/wiki/Genehtala%C5%A1_koda" title="Genehtalaš koda – Northern Sami" lang="se" hreflang="se" data-title="Genehtalaš koda" data-language-autonym="Davvisámegiella" data-language-local-name="Northern Sami" class="interlanguage-link-target"><span>Davvisámegiella</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Genetischer_Code" title="Genetischer Code – German" lang="de" hreflang="de" data-title="Genetischer Code" 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/Geneetiline_kood" title="Geneetiline kood – Estonian" lang="et" hreflang="et" data-title="Geneetiline kood" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%93%CE%B5%CE%BD%CE%B5%CF%84%CE%B9%CE%BA%CF%8C%CF%82_%CE%BA%CF%8E%CE%B4%CE%B9%CE%BA%CE%B1%CF%82" title="Γενετικός κώδικας – Greek" lang="el" hreflang="el" data-title="Γενετικός κώδικας" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/C%C3%B3digo_gen%C3%A9tico" title="Código genético – Spanish" lang="es" hreflang="es" data-title="Código genético" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Genetika_kodo" title="Genetika kodo – Esperanto" lang="eo" hreflang="eo" data-title="Genetika kodo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Kode_genetiko" title="Kode genetiko – Basque" lang="eu" hreflang="eu" data-title="Kode genetiko" 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%B1%D9%85%D8%B2_%DA%98%D9%86%D8%AA%DB%8C%DA%A9%DB%8C" title="رمز ژنتیکی – Persian" lang="fa" hreflang="fa" data-title="رمز ژنتیکی" 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/Code_g%C3%A9n%C3%A9tique" title="Code génétique – French" lang="fr" hreflang="fr" data-title="Code génétique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/C%C3%B3d_g%C3%A9initeach" title="Cód géiniteach – Irish" lang="ga" hreflang="ga" data-title="Cód géiniteach" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/C%C3%B3digo_xen%C3%A9tico" title="Código xenético – Galician" lang="gl" hreflang="gl" data-title="Código xenético" 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/%EC%9C%A0%EC%A0%84_%EB%B6%80%ED%98%B8" title="유전 부호 – Korean" lang="ko" hreflang="ko" data-title="유전 부호" 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/%D4%B3%D5%A5%D5%B6%D5%A5%D5%BF%D5%AB%D5%AF%D5%A1%D5%AF%D5%A1%D5%B6_%D5%AF%D5%B8%D5%A4" title="Գենետիկական կոդ – Armenian" lang="hy" hreflang="hy" data-title="Գենետիկական կոդ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%85%E0%A4%A8%E0%A5%81%E0%A4%B5%E0%A4%BE%E0%A4%82%E0%A4%B6%E0%A4%BF%E0%A4%95_%E0%A4%95%E0%A5%82%E0%A4%9F" title="अनुवांशिक कूट – Hindi" lang="hi" hreflang="hi" data-title="अनुवांशिक कूट" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Genska_uputa" title="Genska uputa – Croatian" lang="hr" hreflang="hr" data-title="Genska uputa" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Codice_genetico" title="Codice genetico – Italian" lang="it" hreflang="it" data-title="Codice genetico" 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%A7%D7%95%D7%93_%D7%94%D7%92%D7%A0%D7%98%D7%99" title="הקוד הגנטי – Hebrew" lang="he" hreflang="he" data-title="הקוד הגנטי" 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%92%E1%83%94%E1%83%9C%E1%83%94%E1%83%A2%E1%83%98%E1%83%99%E1%83%A3%E1%83%A0%E1%83%98_%E1%83%99%E1%83%9D%E1%83%93%E1%83%98" title="გენეტიკური კოდი – Georgian" lang="ka" hreflang="ka" data-title="გენეტიკური კოდი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0%D0%BB%D1%8B%D2%9B_%D0%BA%D0%BE%D0%B4" title="Генетикалық код – Kazakh" lang="kk" hreflang="kk" data-title="Генетикалық код" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Koda_bomawey%C3%AA" title="Koda bomaweyê – Kurdish" lang="ku" hreflang="ku" data-title="Koda bomaweyê" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0%D0%BB%D1%8B%D0%BA_%D0%BA%D0%BE%D0%B4" title="Генетикалык код – Kyrgyz" lang="ky" hreflang="ky" data-title="Генетикалык код" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Codex_geneticus" title="Codex geneticus – Latin" lang="la" hreflang="la" data-title="Codex geneticus" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Genetinis_kodas" title="Genetinis kodas – Lithuanian" lang="lt" hreflang="lt" data-title="Genetinis kodas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Genetikai_k%C3%B3d" title="Genetikai kód – Hungarian" lang="hu" hreflang="hu" data-title="Genetikai kód" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D1%81%D0%BA%D0%B8_%D0%BA%D0%BE%D0%B4" title="Генетски код – Macedonian" lang="mk" hreflang="mk" data-title="Генетски код" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA_%D0%BA%D0%BE%D0%B4" title="Генетик код – Mongolian" lang="mn" hreflang="mn" data-title="Генетик код" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Genetische_code" title="Genetische code – Dutch" lang="nl" hreflang="nl" data-title="Genetische code" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E9%81%BA%E4%BC%9D%E6%83%85%E5%A0%B1" title="遺伝情報 – Japanese" lang="ja" hreflang="ja" data-title="遺伝情報" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Genetisk_kode" title="Genetisk kode – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Genetisk kode" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Den_genetiske_koden" title="Den genetiske koden – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Den genetiske koden" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/C%C3%B2di_genetic" title="Còdi genetic – Occitan" lang="oc" hreflang="oc" data-title="Còdi genetic" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Genetik_kod" title="Genetik kod – Uzbek" lang="uz" hreflang="uz" data-title="Genetik kod" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%A7%D8%B1%D8%AB%D9%8A(%D8%AC%D9%86%DB%8C%D8%AA%DB%8C%DA%A9)_%DA%A9%D9%88%DA%89%D9%88%D9%86%D9%87" title="ارثي(جنیتیک) کوډونه – Pashto" lang="ps" hreflang="ps" data-title="ارثي(جنیتیک) کوډونه" 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/Kod_genetyczny" title="Kod genetyczny – Polish" lang="pl" hreflang="pl" data-title="Kod genetyczny" 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/C%C3%B3digo_gen%C3%A9tico" title="Código genético – Portuguese" lang="pt" hreflang="pt" data-title="Código genético" 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/Cod_genetic" title="Cod genetic – Romanian" lang="ro" hreflang="ro" data-title="Cod genetic" 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-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B8%D0%B9_%D0%BA%D0%BE%D0%B4" title="Генетический код – Russian" lang="ru" hreflang="ru" data-title="Генетический код" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Kodi_gjenetik" title="Kodi gjenetik – Albanian" lang="sq" hreflang="sq" data-title="Kodi gjenetik" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-si badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://si.wikipedia.org/wiki/%E0%B6%A2%E0%B7%8F%E0%B6%B1%E0%B6%B8%E0%B6%BA_%E0%B6%9A%E0%B7%9A%E0%B6%AD" title="ජානමය කේත – Sinhala" lang="si" hreflang="si" data-title="ජානමය කේත" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Genetic_code" title="Genetic code – Simple English" lang="en-simple" hreflang="en-simple" data-title="Genetic code" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Genetick%C3%BD_k%C3%B3d" title="Genetický kód – Slovak" lang="sk" hreflang="sk" data-title="Genetický kód" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Genetski_kod" title="Genetski kod – Slovenian" lang="sl" hreflang="sl" data-title="Genetski kod" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%A8%DB%86%D9%85%D8%A7%D9%88%DB%95_%D9%BE%DB%95%DA%95%D9%84%DB%95" title="بۆماوە پەڕلە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="بۆماوە پەڕلە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D1%87%D0%BA%D0%B8_%D0%BA%D0%BE%D0%B4" title="Генетички код – Serbian" lang="sr" hreflang="sr" data-title="Генетички код" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Genetski_kod" title="Genetski kod – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Genetski kod" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Geneettinen_koodi" title="Geneettinen koodi – Finnish" lang="fi" hreflang="fi" data-title="Geneettinen koodi" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Genetiska_koden" title="Genetiska koden – Swedish" lang="sv" hreflang="sv" data-title="Genetiska koden" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AE%E0%AE%B0%E0%AE%AA%E0%AF%81%E0%AE%95%E0%AF%8D%E0%AE%95%E0%AF%81%E0%AE%B1%E0%AE%BF%E0%AE%AF%E0%AF%80%E0%AE%9F%E0%AF%81" title="மரபுக்குறியீடு – Tamil" lang="ta" hreflang="ta" data-title="மரபுக்குறியீடு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A3%E0%B8%AB%E0%B8%B1%E0%B8%AA%E0%B8%9E%E0%B8%B1%E0%B8%99%E0%B8%98%E0%B8%B8%E0%B8%81%E0%B8%A3%E0%B8%A3%E0%B8%A1" title="รหัสพันธุกรรม – Thai" lang="th" hreflang="th" data-title="รหัสพันธุกรรม" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Genetik_kod" title="Genetik kod – Turkish" lang="tr" hreflang="tr" data-title="Genetik kod" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D1%87%D0%BD%D0%B8%D0%B9_%D0%BA%D0%BE%D0%B4" title="Генетичний код – Ukrainian" lang="uk" hreflang="uk" data-title="Генетичний код" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D9%88%D8%B1%D8%A7%D8%AB%DB%8C_%D8%B1%D9%85%D9%88%D8%B2" title="وراثی رموز – Urdu" lang="ur" hreflang="ur" data-title="وراثی رموز" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://vi.wikipedia.org/wiki/M%C3%A3_di_truy%E1%BB%81n" title="Mã di truyền – Vietnamese" lang="vi" hreflang="vi" data-title="Mã di truyền" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E9%81%97%E4%BC%A0%E5%AF%86%E7%A0%81" title="遗传密码 – Wu" lang="wuu" hreflang="wuu" data-title="遗传密码" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E9%81%97%E4%BC%A0%E5%AF%86%E7%A0%81" title="遗传密码 – Chinese" lang="zh" hreflang="zh" data-title="遗传密码" 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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"><span class="mw-redirectedfrom">(Redirected from <a href="/w/index.php?title=Codon&redirect=no" class="mw-redirect" title="Codon">Codon</a>)</span></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">Rules by which information encoded within genetic material is translated into proteins</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:RNA-codons.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/RNA-codons.svg/220px-RNA-codons.svg.png" decoding="async" width="220" height="311" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/RNA-codons.svg/330px-RNA-codons.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/RNA-codons.svg/440px-RNA-codons.svg.png 2x" data-file-width="744" data-file-height="1052" /></a><figcaption>A series of codons in part of a <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> (mRNA) molecule. Each codon consists of three <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a>, usually corresponding to a single <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a>. The nucleotides are abbreviated with the letters A, U, G and C. This is mRNA, which uses U (<a href="/wiki/Uracil" title="Uracil">uracil</a>). DNA uses T (<a href="/wiki/Thymine" title="Thymine">thymine</a>) instead. This mRNA molecule will instruct a <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> to synthesize a protein according to this code.</figcaption></figure> <p>The <b>genetic code</b> is the set of rules used by living <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a> to <a href="/wiki/Translation_(biology)" title="Translation (biology)">translate</a> information encoded within genetic material (<a href="/wiki/DNA" title="DNA">DNA</a> or <a href="/wiki/RNA" title="RNA">RNA</a> sequences of nucleotide triplets or <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>) into <a href="/wiki/Protein" title="Protein">proteins</a>. Translation is accomplished by the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>, which links <a href="/wiki/Proteinogenic_amino_acid" title="Proteinogenic amino acid">proteinogenic amino acids</a> in an order specified by <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> (mRNA), using <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNA</a> (tRNA) molecules to carry amino acids and to read the mRNA three <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a> at a time. The genetic code is highly similar among all organisms and can be expressed in a simple table with 64 entries. </p><p>The codons specify which amino acid will be added next during <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein biosynthesis</a>. With some exceptions,<sup id="cite_ref-pmid19131629_1-0" class="reference"><a href="#cite_note-pmid19131629-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> a three-nucleotide codon in a nucleic acid sequence specifies a single amino acid. The vast majority of <a href="/wiki/Gene" title="Gene">genes</a> are encoded with a single scheme (see the <a href="/wiki/Codon_tables" class="mw-redirect" title="Codon tables">RNA codon table</a>). That scheme is often called the canonical or standard genetic code, or simply <i>the</i> genetic code, though <a href="#Variations">variant codes</a> (such as in <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a>) exist. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=1" title="Edit section: History"><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:GeneticCode21-version-2.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/GeneticCode21-version-2.svg/330px-GeneticCode21-version-2.svg.png" decoding="async" width="330" height="427" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/GeneticCode21-version-2.svg/495px-GeneticCode21-version-2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/GeneticCode21-version-2.svg/660px-GeneticCode21-version-2.svg.png 2x" data-file-width="579" data-file-height="749" /></a><figcaption>The genetic code</figcaption></figure><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">Further information: <a href="/wiki/Adaptor_hypothesis" title="Adaptor hypothesis">Adaptor hypothesis</a></div> <p>Efforts to understand how proteins are encoded began after <a href="/wiki/Nucleic_acid_double_helix" title="Nucleic acid double helix">DNA's structure</a> was discovered in 1953. The key discoverers, English biophysicist <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> and American biologist <a href="/wiki/James_Watson" title="James Watson">James Watson</a>, working together at the <a href="/wiki/Cavendish_Laboratory" title="Cavendish Laboratory">Cavendish Laboratory</a> of the University of Cambridge, hypothesied that information flows from DNA and that there is a link between DNA and proteins.<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> Soviet-American physicist <a href="/wiki/George_Gamow" title="George Gamow">George Gamow</a> was the first to give a workable scheme for protein synthesis from DNA.<sup id="cite_ref-:3_3-0" class="reference"><a href="#cite_note-:3-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> He postulated that sets of three bases (triplets) must be employed to encode the 20 standard amino acids used by living cells to build proteins, which would allow a maximum of <span class="nowrap">4<sup><small>3</small></sup> = 64</span> amino acids.<sup id="cite_ref-isbn0-465-09138-5_4-0" class="reference"><a href="#cite_note-isbn0-465-09138-5-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> He named this DNA–protein interaction (the original genetic code) as the "diamond code".<sup id="cite_ref-:6_5-0" class="reference"><a href="#cite_note-:6-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>In 1954, Gamow created an informal scientific organisation the <a href="/wiki/RNA_Tie_Club" title="RNA Tie Club">RNA Tie Club</a>, as suggested by Watson, for scientists of different persuasions who were interested in how <a href="/wiki/Translation_(biology)" title="Translation (biology)">proteins were synthesised</a> from genes. However, the club could have only 20 permanent members to represent each of the 20 amino acids; and four additional honorary members to represent the four nucleotides of DNA.<sup id="cite_ref-:5_6-0" class="reference"><a href="#cite_note-:5-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The first scientific contribution of the club, later recorded as "one of the most important unpublished articles in the history of science"<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> and "the most famous unpublished paper in the annals of molecular biology",<sup id="cite_ref-auto_8-0" class="reference"><a href="#cite_note-auto-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> was made by Crick. Crick presented a type-written paper titled "On Degenerate Templates and the Adaptor Hypothesis: A Note for the RNA Tie Club"<sup id="cite_ref-:02_9-0" class="reference"><a href="#cite_note-:02-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> to the members of the club in January 1955, which "totally changed the way we thought about protein synthesis", as Watson recalled.<sup id="cite_ref-:0_10-0" class="reference"><a href="#cite_note-:0-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The hypothesis states that the triplet code was not passed on to amino acids as Gamow thought, but carried by a different molecule, an adaptor, that interacts with amino acids.<sup id="cite_ref-auto_8-1" class="reference"><a href="#cite_note-auto-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The adaptor was later identified as tRNA.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Codons">Codons</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=2" title="Edit section: Codons"><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">"Codon" redirects here. For other uses, see <a href="/wiki/Codon_(disambiguation)" class="mw-disambig" title="Codon (disambiguation)">Codon (disambiguation)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/DNA_and_RNA_codon_tables#Translation_table_1" title="DNA and RNA codon tables">DNA and RNA codon tables § Translation table 1</a></div> <p>The <a href="/wiki/Crick,_Brenner_et_al._experiment" title="Crick, Brenner et al. experiment">Crick, Brenner, Barnett and Watts-Tobin experiment</a> first demonstrated that <b>codons</b> consist of three DNA bases. </p><p><a href="/wiki/Marshall_Nirenberg" class="mw-redirect" title="Marshall Nirenberg">Marshall Nirenberg</a> and <a href="/wiki/J._Heinrich_Matthaei" title="J. Heinrich Matthaei">J. Heinrich Matthaei</a> were the first to reveal the nature of a codon in 1961.<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> They used a <a href="/wiki/Cell-free_system" title="Cell-free system">cell-free system</a> to <a href="/wiki/Translation_(biology)" title="Translation (biology)">translate</a> a poly-<a href="/wiki/Uracil" title="Uracil">uracil</a> RNA sequence (i.e., UUUUU...) and discovered that the <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptide</a> that they had synthesized consisted of only the amino acid <a href="/wiki/Phenylalanine" title="Phenylalanine">phenylalanine</a>.<sup id="cite_ref-pmid14479932_13-0" class="reference"><a href="#cite_note-pmid14479932-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> They thereby deduced that the codon UUU specified the amino acid phenylalanine. </p><p>This was followed by experiments in <a href="/wiki/Severo_Ochoa" title="Severo Ochoa">Severo Ochoa</a>'s laboratory that demonstrated that the poly-<a href="/wiki/Adenine" title="Adenine">adenine</a> RNA sequence (AAAAA...) coded for the polypeptide poly-<a href="/wiki/Lysine" title="Lysine">lysine</a><sup id="cite_ref-pmid13946552_14-0" class="reference"><a href="#cite_note-pmid13946552-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> and that the poly-<a href="/wiki/Cytosine" title="Cytosine">cytosine</a> RNA sequence (CCCCC...) coded for the polypeptide poly-<a href="/wiki/Proline" title="Proline">proline</a>.<sup id="cite_ref-pmid13998282_15-0" class="reference"><a href="#cite_note-pmid13998282-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Therefore, the codon AAA specified the amino acid <a href="/wiki/Lysine" title="Lysine">lysine</a>, and the codon CCC specified the amino acid <a href="/wiki/Proline" title="Proline">proline</a>. Using various <a href="/wiki/Copolymers" class="mw-redirect" title="Copolymers">copolymers</a> most of the remaining codons were then determined. </p><p>Subsequent work by <a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">Har Gobind Khorana</a> identified the rest of the genetic code. Shortly thereafter, <a href="/wiki/Robert_W._Holley" title="Robert W. Holley">Robert W. Holley</a> determined the structure of <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNA</a> (tRNA), the adapter molecule that facilitates the process of translating RNA into protein. This work was based upon Ochoa's earlier studies, yielding the latter the <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> in 1959 for work on the <a href="/wiki/Enzymology" class="mw-redirect" title="Enzymology">enzymology</a> of RNA synthesis.<sup id="cite_ref-Nobel_1959_16-0" class="reference"><a href="#cite_note-Nobel_1959-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Extending this work, Nirenberg and <a href="/wiki/Philip_Leder" title="Philip Leder">Philip Leder</a> revealed the code's triplet nature and deciphered its codons. In these experiments, various combinations of <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> were passed through a filter that contained <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>, the components of cells that <a href="/wiki/Translation_(biology)" title="Translation (biology)">translate</a> RNA into protein. Unique triplets promoted the binding of specific tRNAs to the ribosome. Leder and Nirenberg were able to determine the sequences of 54 out of 64 codons in their experiments.<sup id="cite_ref-pmid5330357_17-0" class="reference"><a href="#cite_note-pmid5330357-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Khorana, Holley and Nirenberg received the Nobel Prize (1968) for their work.<sup id="cite_ref-Nobel_1968_18-0" class="reference"><a href="#cite_note-Nobel_1968-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>The three stop codons were named by discoverers Richard Epstein and Charles Steinberg. "Amber" was named after their friend Harris Bernstein, whose last name means "amber" in German.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> The other two stop codons were named "ochre" and "opal" in order to keep the "color names" theme. </p> <div class="mw-heading mw-heading3"><h3 id="Expanded_genetic_codes_(synthetic_biology)"><span id="Expanded_genetic_codes_.28synthetic_biology.29"></span>Expanded genetic codes (synthetic biology)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=3" title="Edit section: Expanded genetic codes (synthetic biology)"><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">Main article: <a href="/wiki/Expanded_genetic_code" title="Expanded genetic code">Expanded genetic code</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Nucleic_acid_analogues" class="mw-redirect" title="Nucleic acid analogues">Nucleic acid analogues</a></div> <p>In a broad academic audience, the concept of the evolution of the genetic code from the original and ambiguous genetic code to a well-defined ("frozen") code with the repertoire of 20 (+2) canonical amino acids is widely accepted.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> However, there are different opinions, concepts, approaches and ideas, which is the best way to change it experimentally.<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="are the opinions differing on "which one method is the best to change the experiments"? (February 2025)">clarification needed</span></a></i>]</sup> Even models are proposed that predict "entry points" for synthetic amino acid invasion of the genetic code.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Since 2001, 40 non-natural amino acids have been added into proteins by creating a unique codon (recoding) and a corresponding transfer-RNA:aminoacyl – tRNA-synthetase pair to encode it with diverse physicochemical and biological properties in order to be used as a tool to exploring <a href="/wiki/Protein_structure" title="Protein structure">protein structure</a> and function or to create novel or enhanced proteins.<sup id="cite_ref-XieSchultz2005_22-0" class="reference"><a href="#cite_note-XieSchultz2005-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19318213_23-0" class="reference"><a href="#cite_note-pmid19318213-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>H. Murakami and M. Sisido extended some codons to have four and five bases. <a href="/wiki/Steven_A._Benner" title="Steven A. Benner">Steven A. Benner</a> constructed a functional 65th (<i><a href="/wiki/In_vivo" title="In vivo">in vivo</a></i>) codon.<sup id="cite_ref-isbn0-387-22046-1_24-0" class="reference"><a href="#cite_note-isbn0-387-22046-1-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>In 2015 <a href="/wiki/Nediljko_Budisa" title="Nediljko Budisa">N. Budisa</a>, <a href="/wiki/Dieter_S%C3%B6ll" title="Dieter Söll">D. Söll</a> and co-workers reported the full substitution of all 20,899 <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> residues (UGG codons) with unnatural thienopyrrole-alanine in the genetic code of the <a href="/wiki/Bacteria" title="Bacteria">bacterium</a> <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a></i>.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>In 2016 the first stable semisynthetic organism was created. It was a (single cell) bacterium with two synthetic bases (called X and Y). The bases survived cell division.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>In 2017, researchers in South Korea reported that they had engineered a mouse with an extended genetic code that can produce proteins with unnatural amino acids.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>In May 2019, researchers reported the creation of a new "Syn61" strain of the <i>E. coli</i> bacteria. This strain has a fully <a href="/wiki/Synthetic_biology#Synthetic_life" title="Synthetic biology">synthetic</a> genome that is refactored (all overlaps expanded), recoded (removing the use of three out of 64 codons completely), and further modified to remove the now unnecessary tRNAs and release factors. It is fully <a href="/wiki/Genetic_viability" title="Genetic viability">viable</a> and grows 1.6× slower than its wild-type counterpart "<a href="/wiki/Escherichia_coli#MDS42" title="Escherichia coli">MDS42</a>".<sup id="cite_ref-NYT-20190515_29-0" class="reference"><a href="#cite_note-NYT-20190515-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NAT-20190515_30-0" class="reference"><a href="#cite_note-NAT-20190515-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Features">Features</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=4" title="Edit section: Features"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg/250px-Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg.png" decoding="async" width="220" height="118" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg/330px-Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/af/Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg/500px-Homo_sapiens-mtDNA~NC_012920-ATP8%2BATP6_Overlap.svg.png 2x" data-file-width="716" data-file-height="384" /></a><figcaption>Reading frames in the DNA sequence of a region of the human mitochondrial genome coding for the genes <i><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a></i> and <i><a href="/wiki/MT-ATP6" title="MT-ATP6">MT-ATP6</a></i> (in black: positions 8,525 to 8,580 in the sequence accession NC_012920<sup id="cite_ref-NCBI_NC_012920_31-0" class="reference"><a href="#cite_note-NCBI_NC_012920-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup>). There are three possible reading frames in the 5' → 3' forward direction, starting on the first (+1), second (+2) and third position (+3). For each codon (square brackets), the amino acid is given by the <a href="/wiki/Vertebrate_mitochondrial_code" title="Vertebrate mitochondrial code">vertebrate mitochondrial code</a>, either in the +1 frame for <i>MT-ATP8</i> (in red) or in the +3 frame for <i>MT-ATP6</i> (in blue). The <i>MT-ATP8</i> genes terminates with the TAG stop codon (red dot) in the +1 frame. The <i>MT-ATP6</i> gene starts with the ATG codon (blue circle for the M amino acid) in the +3 frame.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Reading_frame">Reading frame</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=5" title="Edit section: Reading frame"><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">Main article: <a href="/wiki/Reading_frame" title="Reading frame">Reading frame</a></div> <p>A reading frame is defined by the initial triplet of nucleotides from which translation starts. It sets the frame for a run of successive, non-overlapping codons, which is known as an "<a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frame</a>" (ORF). For example, the string 5'-AAATGAACG-3' (see figure), if read from the first position, contains the codons AAA, TGA, and ACG ; if read from the second position, it contains the codons AAT and GAA ; and if read from the third position, it contains the codons ATG and AAC. Every sequence can, thus, be read in its <a href="/wiki/5%27_to_3%27" class="mw-redirect" title="5' to 3'">5' → 3' direction</a> in three <a href="/wiki/Reading_frames" class="mw-redirect" title="Reading frames">reading frames</a>, each producing a possibly distinct amino acid sequence: in the given example, Lys (K)-Trp (W)-Thr (T), Asn (N)-Glu (E), or Met (M)-Asn (N), respectively (when translating with the <a href="/wiki/Vertebrate_mitochondrial_code" title="Vertebrate mitochondrial code">vertebrate mitochondrial code</a>). When DNA is double-stranded, six possible <a href="/wiki/Reading_frames" class="mw-redirect" title="Reading frames">reading frames</a> are defined, three in the forward orientation on one strand and three reverse on the opposite strand.<sup id="cite_ref-genetics_dictionary_32-0" class="reference"><a href="#cite_note-genetics_dictionary-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 330">: 330 </span></sup> Protein-coding frames are defined by a <a href="/wiki/Start_codon" title="Start codon">start codon</a>, usually the first AUG codon in the RNA, (ATG in DNA) sequence. </p><p>In <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, open reading frames in <a href="/wiki/Exon" title="Exon">exons</a> are often interrupted by <a href="/wiki/Intron" title="Intron">introns</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Start_and_stop_codons">Start and stop codons</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=6" title="Edit section: Start and stop codons"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Translation starts with a chain-initiation codon or <a href="/wiki/Start_codon" title="Start codon">start codon</a>. The start codon alone is not sufficient to begin the process. Nearby sequences such as the <a href="/wiki/Shine-Dalgarno" class="mw-redirect" title="Shine-Dalgarno">Shine-Dalgarno</a> sequence in <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a></i> and <a href="/wiki/Initiation_factor" title="Initiation factor">initiation factors</a> are also required to start translation. The most common start codon is AUG, which is read as <a href="/wiki/Methionine" title="Methionine">methionine</a> or as <a href="/wiki/N-Formylmethionine" title="N-Formylmethionine">formylmethionine</a> (in bacteria, mitochondria, and plastids). Alternative start codons depending on the organism include "GUG" or "UUG"; these codons normally represent <a href="/wiki/Valine" title="Valine">valine</a> and <a href="/wiki/Leucine" title="Leucine">leucine</a>, respectively, but as start codons they are translated as methionine or formylmethionine.<sup id="cite_ref-pmid12867081_33-0" class="reference"><a href="#cite_note-pmid12867081-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p><p>The three <a href="/wiki/Stop_codon" title="Stop codon">stop codons</a> have names: UAG is <i>amber</i>, UGA is <i>opal</i> (sometimes also called <i>umber</i>), and UAA is <i>ochre</i>. Stop codons are also called "termination" or "nonsense" codons. They signal release of the nascent polypeptide from the ribosome because no cognate tRNA has anticodons complementary to these stop signals, allowing a <a href="/wiki/Release_factor" title="Release factor">release factor</a> to bind to the ribosome instead.<sup id="cite_ref-urlHow_nonsense_mutations_got_their_names_34-0" class="reference"><a href="#cite_note-urlHow_nonsense_mutations_got_their_names-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Effect_of_mutations">Effect of mutations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=7" title="Edit section: Effect of mutations"><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:Notable_mutations.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Notable_mutations.svg/500px-Notable_mutations.svg.png" decoding="async" width="390" height="231" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Notable_mutations.svg/960px-Notable_mutations.svg.png 1.5x" data-file-width="2980" data-file-height="1764" /></a><figcaption>Examples of notable <a href="/wiki/Mutation" title="Mutation">mutations</a> that can occur in humans<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>During the process of <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>, errors occasionally occur in the <a href="/wiki/Polymerization" title="Polymerization">polymerization</a> of the second strand. These errors, <a href="/wiki/Mutation" title="Mutation">mutations</a>, can affect an organism's <a href="/wiki/Phenotype" title="Phenotype">phenotype</a>, especially if they occur within the protein coding sequence of a gene. Error rates are typically 1 error in every 10–100 million bases—due to the "<a href="/wiki/Proofreading_(biology)" title="Proofreading (biology)">proofreading</a>" ability of <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerases</a>.<sup id="cite_ref-griffiths2000sect2706_36-0" class="reference"><a href="#cite_note-griffiths2000sect2706-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kunkel_37-0" class="reference"><a href="#cite_note-Kunkel-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Missense_mutation" title="Missense mutation">Missense mutations</a> and <a href="/wiki/Nonsense_mutation" title="Nonsense mutation">nonsense mutations</a> are examples of <a href="/wiki/Point_mutation" title="Point mutation">point mutations</a> that can cause genetic diseases such as <a href="/wiki/Sickle-cell_disease" class="mw-redirect" title="Sickle-cell disease">sickle-cell disease</a> and <a href="/wiki/Thalassemia" title="Thalassemia">thalassemia</a> respectively.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid88735_39-0" class="reference"><a href="#cite_note-pmid88735-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid17015226_40-0" class="reference"><a href="#cite_note-pmid17015226-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Clinically important missense mutations generally change the properties of the coded amino acid residue among basic, acidic, polar or non-polar states, whereas nonsense mutations result in a <a href="/wiki/Stop_codon" title="Stop codon">stop codon</a>.<sup id="cite_ref-genetics_dictionary_32-1" class="reference"><a href="#cite_note-genetics_dictionary-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Mutations that disrupt the reading frame sequence by <a href="/wiki/Indels" class="mw-redirect" title="Indels">indels</a> (<a href="/wiki/Gene_insertion" class="mw-redirect" title="Gene insertion">insertions</a> or <a href="/wiki/Genetic_deletion" class="mw-redirect" title="Genetic deletion">deletions</a>) of a non-multiple of 3 nucleotide bases are known as <a href="/wiki/Frameshift_mutation" title="Frameshift mutation">frameshift mutations</a>. These mutations usually result in a completely different translation from the original, and likely cause a <a href="/wiki/Stop_codon" title="Stop codon">stop codon</a> to be read, which truncates the protein.<sup id="cite_ref-pmid8723688_41-0" class="reference"><a href="#cite_note-pmid8723688-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> These mutations may impair the protein's function and are thus rare in <i><a href="/wiki/In_vivo" title="In vivo">in vivo</a></i> protein-coding sequences. One reason inheritance of frameshift mutations is rare is that, if the protein being translated is essential for growth under the selective pressures the organism faces, absence of a functional protein may cause death before the organism becomes viable.<sup id="cite_ref-pmid8444142_42-0" class="reference"><a href="#cite_note-pmid8444142-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Frameshift mutations may result in severe genetic diseases such as <a href="/wiki/Tay%E2%80%93Sachs_disease" title="Tay–Sachs disease">Tay–Sachs disease</a>.<sup id="cite_ref-isbn0-07-111156-5_43-0" class="reference"><a href="#cite_note-isbn0-07-111156-5-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>Although most mutations that change protein sequences are harmful or neutral, some mutations have benefits.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> These mutations may enable the mutant organism to withstand particular environmental stresses better than <a href="/wiki/Wild_type" title="Wild type">wild type</a> organisms, or reproduce more quickly. In these cases a mutation will tend to become more common in a population through <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Virus" title="Virus">Viruses</a> that use <a href="/wiki/RNA" title="RNA">RNA</a> as their genetic material have rapid mutation rates,<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> which can be an advantage, since these viruses thereby evolve rapidly, and thus evade the <a href="/wiki/Immune_system" title="Immune system">immune system</a> defensive responses.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> In large populations of asexually reproducing organisms, for example, <i>E. coli</i>, multiple beneficial mutations may co-occur. This phenomenon is called <a href="/wiki/Clonal_interference" title="Clonal interference">clonal interference</a> and causes competition among the mutations.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Degeneracy">Degeneracy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=8" title="Edit section: Degeneracy"><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">Main article: <a href="/wiki/Codon_degeneracy" title="Codon degeneracy">Codon degeneracy</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Genetic_Code_Simple_Corrected.pdf" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Genetic_Code_Simple_Corrected.pdf/page1-220px-Genetic_Code_Simple_Corrected.pdf.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Genetic_Code_Simple_Corrected.pdf/page1-330px-Genetic_Code_Simple_Corrected.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/84/Genetic_Code_Simple_Corrected.pdf/page1-440px-Genetic_Code_Simple_Corrected.pdf.jpg 2x" data-file-width="1350" data-file-height="1350" /></a><figcaption>Grouping of codons by amino acid residue molar volume and <a href="/wiki/Hydropathicity" class="mw-redirect" title="Hydropathicity">hydropathicity</a>. A <a href="/wiki/File:ELLIPTICAL_GENETIC_CODE_Ian.png" title="File:ELLIPTICAL GENETIC CODE Ian.png">more detailed version</a> is available.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:3D_Genetic_Code.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/74/3D_Genetic_Code.jpg/250px-3D_Genetic_Code.jpg" decoding="async" width="220" height="229" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/74/3D_Genetic_Code.jpg/330px-3D_Genetic_Code.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/74/3D_Genetic_Code.jpg/500px-3D_Genetic_Code.jpg 2x" data-file-width="1291" data-file-height="1345" /></a><figcaption>Axes 1, 2, 3 are the first, second, and third positions in the codon. The 20 amino acids and stop codons (X) are shown in <a href="/wiki/Amino_acid#Table_of_standard_amino_acid_abbreviations_and_properties" title="Amino acid">single letter code</a>.</figcaption></figure> <p>Degeneracy is the redundancy of the genetic code. This term was given by Bernfield and Nirenberg. The genetic code has redundancy but no ambiguity (see the <a href="/wiki/DNA_and_RNA_codon_tables" title="DNA and RNA codon tables">codon tables</a> below for the full correlation). For example, although codons GAA and GAG both specify <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamic acid</a> (redundancy), neither specifies another amino acid (no ambiguity). The codons encoding one amino acid may differ in any of their three positions. For example, the amino acid leucine is specified by <b>Y</b>U<b>R</b> or CU<b>N</b> (UUA, UUG, CUU, CUC, CUA, or CUG) codons (difference in the first or third position indicated using <a href="/wiki/Nucleic_acid_notation" title="Nucleic acid notation">IUPAC notation</a>), while the amino acid <a href="/wiki/Serine" title="Serine">serine</a> is specified by UC<b>N</b> or AG<b>Y</b> (UCA, UCG, UCC, UCU, AGU, or AGC) codons (difference in the first, second, or third position).<sup id="cite_ref-MBG_49-0" class="reference"><a href="#cite_note-MBG-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> A practical consequence of redundancy is that errors in the third position of the triplet codon cause only a silent mutation or an error that would not affect the protein because the <a href="/wiki/Hydrophilicity" class="mw-redirect" title="Hydrophilicity">hydrophilicity</a> or <a href="/wiki/Hydrophobicity" class="mw-redirect" title="Hydrophobicity">hydrophobicity</a> is maintained by equivalent substitution of amino acids; for example, a codon of NUN (where N = any nucleotide) tends to code for hydrophobic amino acids. NCN yields amino acid residues that are small in size and moderate in <a href="/wiki/Hydropathicity" class="mw-redirect" title="Hydropathicity">hydropathicity</a>; NAN encodes average size hydrophilic residues. The genetic code is so well-structured for hydropathicity that a mathematical analysis (<a href="/wiki/Singular_value_decomposition" title="Singular value decomposition">Singular Value Decomposition</a>) of 12 variables (4 nucleotides x 3 positions) yields a remarkable correlation (C = 0.95) for predicting the hydropathicity of the encoded amino acid directly from the triplet nucleotide sequence, <i>without translation.</i><sup id="cite_ref-Michel-Beyerle1990_50-0" class="reference"><a href="#cite_note-Michel-Beyerle1990-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> Note in the table, below, eight amino acids are not affected at all by mutations at the third position of the codon, whereas in the figure above, a mutation at the second position is likely to cause a radical change in the physicochemical properties of the encoded amino acid. Nevertheless, changes in the first position of the codons are more important than changes in the second position on a global scale.<sup id="cite_ref-Fricke_52-0" class="reference"><a href="#cite_note-Fricke-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> The reason may be that charge reversal (from a positive to a negative charge or vice versa) can only occur upon mutations in the first position of certain codons, but not upon changes in the second position of any codon. Such charge reversal may have dramatic consequences for the structure or function of a protein. This aspect may have been largely underestimated by previous studies.<sup id="cite_ref-Fricke_52-1" class="reference"><a href="#cite_note-Fricke-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Codon_usage_bias">Codon usage bias</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=9" title="Edit section: Codon usage bias"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The frequency of codons, also known as <a href="/wiki/Codon_usage_bias" title="Codon usage bias">codon usage bias</a>, can vary from species to species with functional implications for the control of <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>. The codon varies by organism; for example, most common proline codon in E. coli is CCG, whereas in humans this is the least used proline codon.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1256386598">.mw-parser-output .cot-header-mainspace{background:#F0F2F5;color:inherit}.mw-parser-output .cot-header-other{background:#CCFFCC;color:inherit}@media screen{html.skin-theme-clientpref-night .mw-parser-output .cot-header-mainspace{background:#14181F;color:inherit}html.skin-theme-clientpref-night .mw-parser-output .cot-header-other{background:#003500;color:inherit}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cot-header-mainspace{background:#14181F;color:inherit}html.skin-theme-clientpref-os .mw-parser-output .cot-header-other{background:#003500;color:inherit}}</style> <div style="margin-left:0"> <table class="mw-collapsible mw-archivedtalk mw-collapsed" style="color:inherit; background: transparent; text-align: left; border: 1px solid Silver; margin: 0.2em auto auto; width:100%; clear: both; padding: 1px;"> <tbody><tr> <th class="cot-header-mainspace" style="; font-size:87%; padding:0.2em 0.3em; text-align:center;"><div style="font-size:115%;margin:0 4em">Human genome codon frequency table<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup></div> </th></tr> <tr> <td style="color:inherit; border: solid 1px Silver; padding: 0.6em; background: var(--background-color-base, #fff);"> <table class="wikitable" style="text-align: center;"> <tbody><tr> <th>Codon</th> <th>AA<style data-mw-deduplicate="TemplateStyles:r1041539562">.mw-parser-output .citation{word-wrap:break-word}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}</style><sup class="citation nobold" id="ref_startC"><a href="#endnote_startC">[C]</a></sup></th> <th>Fraction<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1041539562" /><sup class="citation nobold" id="ref_startD"><a href="#endnote_startD">[D]</a></sup></th> <th>Freq <a href="/wiki/Per_thousand" class="mw-redirect" title="Per thousand">‰</a><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1041539562" /><sup class="citation nobold" id="ref_startE"><a href="#endnote_startE">[E]</a></sup></th> <th>Number<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1041539562" /><sup class="citation nobold" id="ref_startF"><a href="#endnote_startF">[F]</a></sup> </th> <td rowspan="17"> </td> <th>Codon</th> <th>AA</th> <th>Fraction</th> <th>Freq <a href="/wiki/Per_thousand" class="mw-redirect" title="Per thousand">‰</a></th> <th>Number </th> <td rowspan="17"> </td> <th>Codon</th> <th>AA</th> <th>Fraction</th> <th>Freq <a href="/wiki/Per_thousand" class="mw-redirect" title="Per thousand">‰</a></th> <th>Number </th> <td rowspan="17"> </td> <th>Codon</th> <th>AA</th> <th>Fraction</th> <th>Freq <a href="/wiki/Per_thousand" class="mw-redirect" title="Per thousand">‰</a></th> <th>Number </th></tr> <tr> <td>UUU</td> <td>F</td> <td>0.46</td> <td>17.6</td> <td>714,298</td> <td>UCU</td> <td>S</td> <td>0.19</td> <td>15.2</td> <td>618,711</td> <td>UAU</td> <td>Y</td> <td>0.44</td> <td>12.2</td> <td>495,699</td> <td>UGU</td> <td>C</td> <td>0.46</td> <td>10.6</td> <td>430,311 </td></tr> <tr> <td>UUC</td> <td>F</td> <td>0.54</td> <td>20.3</td> <td>824,692</td> <td>UCC</td> <td>S</td> <td>0.22</td> <td>17.7</td> <td>718,892</td> <td>UAC</td> <td>Y</td> <td>0.56</td> <td>15.3</td> <td>622,407</td> <td>UGC</td> <td>C</td> <td>0.54</td> <td>12.6</td> <td>513,028 </td></tr> <tr> <td>UUA</td> <td>L</td> <td>0.08</td> <td>7.7</td> <td>311,881</td> <td>UCA</td> <td>S</td> <td>0.15</td> <td>12.2</td> <td>496,448</td> <td>UAA</td> <td>*</td> <td>0.30</td> <td>1.0</td> <td>40,285</td> <td>UGA</td> <td>*</td> <td>0.47</td> <td>1.6</td> <td>63,237 </td></tr> <tr> <td>UUG</td> <td>L</td> <td>0.13</td> <td>12.9</td> <td>525,688</td> <td>UCG</td> <td>S</td> <td>0.05</td> <td>4.4</td> <td>179,419</td> <td>UAG</td> <td>*</td> <td>0.24</td> <td>0.8</td> <td>32,109</td> <td>UGG</td> <td>W</td> <td>1.00</td> <td>13.2</td> <td>535,595 </td></tr> <tr> <td>CUU</td> <td>L</td> <td>0.13</td> <td>13.2</td> <td>536,515</td> <td>CCU</td> <td>P</td> <td>0.29</td> <td>17.5</td> <td>713,233</td> <td>CAU</td> <td>H</td> <td>0.42</td> <td>10.9</td> <td>441,711</td> <td>CGU</td> <td>R</td> <td>0.08</td> <td>4.5</td> <td>184,609 </td></tr> <tr> <td>CUC</td> <td>L</td> <td>0.20</td> <td>19.6</td> <td>796,638</td> <td>CCC</td> <td>P</td> <td>0.32</td> <td>19.8</td> <td>804,620</td> <td>CAC</td> <td>H</td> <td>0.58</td> <td>15.1</td> <td>613,713</td> <td>CGC</td> <td>R</td> <td>0.18</td> <td>10.4</td> <td>423,516 </td></tr> <tr> <td>CUA</td> <td>L</td> <td>0.07</td> <td>7.2</td> <td>290,751</td> <td>CCA</td> <td>P</td> <td>0.28</td> <td>16.9</td> <td>688,038</td> <td>CAA</td> <td>Q</td> <td>0.27</td> <td>12.3</td> <td>501,911</td> <td>CGA</td> <td>R</td> <td>0.11</td> <td>6.2</td> <td>250,760 </td></tr> <tr> <td>CUG</td> <td>L</td> <td>0.40</td> <td>39.6</td> <td>1,611,801</td> <td>CCG</td> <td>P</td> <td>0.11</td> <td>6.9</td> <td>281,570</td> <td>CAG</td> <td>Q</td> <td>0.73</td> <td>34.2</td> <td>1,391,973</td> <td>CGG</td> <td>R</td> <td>0.20</td> <td>11.4</td> <td>464,485 </td></tr> <tr> <td>AUU</td> <td>I</td> <td>0.36</td> <td>16.0</td> <td>650,473</td> <td>ACU</td> <td>T</td> <td>0.25</td> <td>13.1</td> <td>533,609</td> <td>AAU</td> <td>N</td> <td>0.47</td> <td>17.0</td> <td>689,701</td> <td>AGU</td> <td>S</td> <td>0.15</td> <td>12.1</td> <td>493,429 </td></tr> <tr> <td>AUC</td> <td>I</td> <td>0.47</td> <td>20.8</td> <td>846,466</td> <td>ACC</td> <td>T</td> <td>0.36</td> <td>18.9</td> <td>768,147</td> <td>AAC</td> <td>N</td> <td>0.53</td> <td>19.1</td> <td>776,603</td> <td>AGC</td> <td>S</td> <td>0.24</td> <td>19.5</td> <td>791,383 </td></tr> <tr> <td>AUA</td> <td>I</td> <td>0.17</td> <td>7.5</td> <td>304,565</td> <td>ACA</td> <td>T</td> <td>0.28</td> <td>15.1</td> <td>614,523</td> <td>AAA</td> <td>K</td> <td>0.43</td> <td>24.4</td> <td>993,621</td> <td>AGA</td> <td>R</td> <td>0.21</td> <td>12.2</td> <td>494,682 </td></tr> <tr> <td>AUG</td> <td>M</td> <td>1.00</td> <td>22.0</td> <td>896,005</td> <td>ACG</td> <td>T</td> <td>0.11</td> <td>6.1</td> <td>246,105</td> <td>AAG</td> <td>K</td> <td>0.57</td> <td>31.9</td> <td>1,295,568</td> <td>AGG</td> <td>R</td> <td>0.21</td> <td>12.0</td> <td>486,463 </td></tr> <tr> <td>GUU</td> <td>V</td> <td>0.18</td> <td>11.0</td> <td>448,607</td> <td>GCU</td> <td>A</td> <td>0.27</td> <td>18.4</td> <td>750,096</td> <td>GAU</td> <td>D</td> <td>0.46</td> <td>21.8</td> <td>885,429</td> <td>GGU</td> <td>G</td> <td>0.16</td> <td>10.8</td> <td>437,126 </td></tr> <tr> <td>GUC</td> <td>V</td> <td>0.24</td> <td>14.5</td> <td>588,138</td> <td>GCC</td> <td>A</td> <td>0.40</td> <td>27.7</td> <td>1,127,679</td> <td>GAC</td> <td>D</td> <td>0.54</td> <td>25.1</td> <td>1,020,595</td> <td>GGC</td> <td>G</td> <td>0.34</td> <td>22.2</td> <td>903,565 </td></tr> <tr> <td>GUA</td> <td>V</td> <td>0.12</td> <td>7.1</td> <td>287,712</td> <td>GCA</td> <td>A</td> <td>0.23</td> <td>15.8</td> <td>643,471</td> <td>GAA</td> <td>E</td> <td>0.42</td> <td>29.0</td> <td>1,177,632</td> <td>GGA</td> <td>G</td> <td>0.25</td> <td>16.5</td> <td>669,873 </td></tr> <tr> <td>GUG</td> <td>V</td> <td>0.46</td> <td>28.1</td> <td>1,143,534</td> <td>GCG</td> <td>A</td> <td>0.11</td> <td>7.4</td> <td>299,495</td> <td>GAG</td> <td>E</td> <td>0.58</td> <td>39.6</td> <td>1,609,975</td> <td>GGG</td> <td>G</td> <td>0.25</td> <td>16.5</td> <td>669,768 </td></tr></tbody></table> </td></tr></tbody></table></div> <div class="mw-heading mw-heading2"><h2 id="Alternative_genetic_codes">Alternative genetic codes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=10" title="Edit section: Alternative genetic codes"><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">See also: <a href="/wiki/DNA_and_RNA_codon_tables#Alternative_codons" title="DNA and RNA codon tables">DNA and RNA codon tables § Alternative codons</a></div> <div class="mw-heading mw-heading3"><h3 id="Non-standard_amino_acids">Non-standard amino acids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=11" title="Edit section: Non-standard amino acids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In some proteins, non-standard amino acids are substituted for standard stop codons, depending on associated signal sequences in the messenger RNA. For example, UGA can code for <a href="/wiki/Selenocysteine" title="Selenocysteine">selenocysteine</a> and UAG can code for <a href="/wiki/Pyrrolysine" title="Pyrrolysine">pyrrolysine</a>. Selenocysteine came to be seen as the 21st amino acid, and pyrrolysine as the 22nd.<sup id="cite_ref-Zhang2005_55-0" class="reference"><a href="#cite_note-Zhang2005-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Both selenocysteine and pyrrolysine may be present in the same organism.<sup id="cite_ref-Zhang2005_55-1" class="reference"><a href="#cite_note-Zhang2005-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Although the genetic code is normally fixed in an organism, the achaeal prokaryote <i><a href="/wiki/Acetohalobium_arabaticum" title="Acetohalobium arabaticum">Acetohalobium arabaticum</a></i> can expand its genetic code from 20 to 21 amino acids (by including pyrrolysine) under different conditions of growth.<sup id="cite_ref-Prat2012_56-0" class="reference"><a href="#cite_note-Prat2012-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Variations">Variations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=12" title="Edit section: Variations"><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">See also: <a href="/wiki/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:FACIL_genetic_code_logo.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/FACIL_genetic_code_logo.png/960px-FACIL_genetic_code_logo.png" decoding="async" width="510" height="55" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/FACIL_genetic_code_logo.png/1020px-FACIL_genetic_code_logo.png 2x" data-file-width="5120" data-file-height="550" /></a><figcaption>Genetic code <a href="/wiki/Sequence_logo" title="Sequence logo">logo</a> of the <i>Globobulimina pseudospinescens</i> mitochondrial genome by FACIL. The program is able to correctly infer that the <a href="/wiki/The_mold,_protozoan,_and_coelenterate_mitochondrial_code_and_the_mycoplasma/spiroplasma_code" title="The mold, protozoan, and coelenterate mitochondrial code and the mycoplasma/spiroplasma code">Protozoan Mitochondrial Code</a> is in use.<sup id="cite_ref-DutilhJurgelenaite2011_57-0" class="reference"><a href="#cite_note-DutilhJurgelenaite2011-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> The logo shows the 64 codons from left to right, predicted alternatives in red (relative to the standard genetic code). Red line: stop codons. The height of each amino acid in the stack shows how often it is aligned to the codon in homologous protein domains. The stack height indicates the support for the prediction.</figcaption></figure> <p>There was originally a simple and widely accepted argument that the genetic code should be universal: namely, that any variation in the genetic code would be lethal to the organism (although Crick had stated that viruses were an exception). This is known as the "frozen accident" argument for the universality of the genetic code. However, in his seminal paper on the origins of the genetic code in 1968, Francis Crick still stated that the universality of the genetic code in all organisms was an unproven assumption, and was probably not true in some instances. He predicted that "The code is universal (the same in all organisms) or nearly so".<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> The first variation was discovered in 1979, by researchers studying <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">human mitochondrial genes</a>.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Many slight variants were discovered thereafter,<sup id="cite_ref-url_The_Genetic_Codes_NCBI_60-0" class="reference"><a href="#cite_note-url_The_Genetic_Codes_NCBI-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> including various alternative mitochondrial codes.<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> These minor variants for example involve translation of the codon UGA as <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> in <i><a href="/wiki/Mycoplasma" title="Mycoplasma">Mycoplasma</a></i> species, and translation of CUG as a serine rather than leucine in yeasts of the "CTG clade" (such as <i><a href="/wiki/Candida_albicans" title="Candida albicans">Candida albicans</a></i>).<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> Because viruses must use the same genetic code as their hosts, modifications to the standard genetic code could interfere with viral protein synthesis or functioning. However, viruses such as <a href="/wiki/Totivirus" title="Totivirus">totiviruses</a> have adapted to the host's genetic code modification.<sup id="cite_ref-pmid23638388_65-0" class="reference"><a href="#cite_note-pmid23638388-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>, GUG and UUG are common start codons. In rare cases, certain proteins may use alternative start codons.<sup id="cite_ref-url_The_Genetic_Codes_NCBI_60-1" class="reference"><a href="#cite_note-url_The_Genetic_Codes_NCBI-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Surprisingly, variations in the interpretation of the genetic code exist also in human nuclear-encoded genes: In 2016, researchers studying the translation of malate dehydrogenase found that in about 4% of the mRNAs encoding this enzyme the stop codon is naturally used to encode the amino acids tryptophan and arginine.<sup id="cite_ref-pmid27881739_66-0" class="reference"><a href="#cite_note-pmid27881739-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> This type of recoding is induced by a high-readthrough stop codon context<sup id="cite_ref-pmid25247702_67-0" class="reference"><a href="#cite_note-pmid25247702-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> and it is referred to as <i>functional translational readthrough</i>.<sup id="cite_ref-PMC4973966_68-0" class="reference"><a href="#cite_note-PMC4973966-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p><p>Despite these differences, all known naturally occurring codes are very similar. The coding mechanism is the same for all organisms: three-base codons, <a href="/wiki/Transfer_RNA" title="Transfer RNA">tRNA</a>, ribosomes, single direction reading and translating single codons into single amino acids.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> The most extreme variations occur in certain ciliates where the meaning of stop codons depends on their position within mRNA. When close to the 3' end they act as terminators while in internal positions they either code for amino acids as in <i><a href="/wiki/Condylostoma" title="Condylostoma">Condylostoma</a> magnum</i><sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> or trigger <a href="/wiki/Ribosomal_frameshift" title="Ribosomal frameshift">ribosomal frameshifting</a> as in <i><a href="/wiki/Euplotes" title="Euplotes">Euplotes</a></i>.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> </p><p>The origins and variation of the genetic code, including the mechanisms behind the evolvability of the genetic code, have been widely studied,<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> and some studies have been done experimentally evolving the genetic code of some organisms.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inference">Inference</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=13" title="Edit section: Inference"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Variant genetic codes used by an organism can be inferred by identifying highly conserved genes encoded in that genome, and comparing its codon usage to the amino acids in homologous proteins of other organisms. For example, the program FACIL infers a genetic code by searching which amino acids in homologous protein domains are most often aligned to every codon. The resulting amino acid (or stop codon) probabilities for each codon are displayed in a genetic code logo.<sup id="cite_ref-DutilhJurgelenaite2011_57-1" class="reference"><a href="#cite_note-DutilhJurgelenaite2011-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>As of January 2022, the most complete survey of genetic codes is done by Shulgina and Eddy, who screened 250,000 prokaryotic genomes using their Codetta tool. This tool uses a similar approach to FACIL with a larger <a href="/wiki/Pfam" title="Pfam">Pfam</a> database. Despite the NCBI already providing 27 translation tables, the authors were able to find new 5 genetic code variations (corroborated by tRNA mutations) and correct several misattributions.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> Codetta was later used to analyze genetic code change in <a href="/wiki/Ciliates" class="mw-redirect" title="Ciliates">ciliates</a>.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Origin">Origin</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=14" title="Edit section: Origin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The genetic code is a key part of the <a href="/wiki/Origin_of_life" class="mw-redirect" title="Origin of life">history of life</a>, according to one version of which self-replicating RNA molecules preceded life as we know it. This is the <a href="/wiki/RNA_world_hypothesis" class="mw-redirect" title="RNA world hypothesis">RNA world hypothesis</a>. Under this hypothesis, any model for the emergence of the genetic code is intimately related to a model of the transfer from <a href="/wiki/Ribozyme" title="Ribozyme">ribozymes</a> (RNA enzymes) to proteins as the principal enzymes in cells. In line with the RNA world hypothesis, transfer RNA molecules appear to have evolved before modern <a href="/wiki/Aminoacyl-tRNA_synthetase" class="mw-redirect" title="Aminoacyl-tRNA synthetase">aminoacyl-tRNA synthetases</a>, so the latter cannot be part of the explanation of its patterns.<sup id="cite_ref-De1998_80-0" class="reference"><a href="#cite_note-De1998-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p><p>A hypothetical randomly evolved genetic code further motivates a biochemical or evolutionary model for its origin. If amino acids were randomly assigned to triplet codons, there would be 1.5 × 10<sup>84</sup> possible genetic codes.<sup id="cite_ref-isbn0-674-05075-4_81-0" class="reference"><a href="#cite_note-isbn0-674-05075-4-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 163">: <a rel="nofollow" class="external text" href="https://books.google.com/books?id=-YLBMmJE1WwC&pg=PA163">163</a> </span></sup> This number is found by calculating the number of ways that 21 items (20 amino acids plus one stop) can be placed in 64 bins, wherein each item is used at least once.<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> However, the distribution of codon assignments in the genetic code is nonrandom.<sup id="cite_ref-pmid9732450_83-0" class="reference"><a href="#cite_note-pmid9732450-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> In particular, the genetic code clusters certain amino acid assignments. </p><p>Amino acids that share the same biosynthetic pathway tend to have the same first base in their codons. This could be an evolutionary relic of an early, simpler genetic code with fewer amino acids that later evolved to code a larger set of amino acids.<sup id="cite_ref-pmid2650752_84-0" class="reference"><a href="#cite_note-pmid2650752-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> It could also reflect steric and chemical properties that had another effect on the codon during its evolution. Amino acids with similar physical properties also tend to have similar codons,<sup id="cite_ref-pmid2514270_85-0" class="reference"><a href="#cite_note-pmid2514270-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid6928661_86-0" class="reference"><a href="#cite_note-pmid6928661-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> reducing the problems caused by point mutations and mistranslations.<sup id="cite_ref-pmid9732450_83-1" class="reference"><a href="#cite_note-pmid9732450-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p><p>Given the non-random genetic triplet coding scheme, a tenable hypothesis for the origin of genetic code could address multiple aspects of the codon table, such as absence of codons for D-amino acids, secondary codon patterns for some amino acids, confinement of synonymous positions to third position, the small set of only 20 amino acids (instead of a number approaching 64), and the relation of stop codon patterns to amino acid coding patterns.<sup id="cite_ref-pmid21779963_87-0" class="reference"><a href="#cite_note-pmid21779963-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p><p>Three main hypotheses address the origin of the genetic code. Many models belong to one of them or to a hybrid:<sup id="cite_ref-pmid10742043_88-0" class="reference"><a href="#cite_note-pmid10742043-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> </p> <ul><li>Random freeze: the genetic code was randomly created. For example, early <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a>-like ribozymes may have had different affinities for amino acids, with codons emerging from another part of the ribozyme that exhibited random variability. Once enough <a href="/wiki/Peptide" title="Peptide">peptides</a> were coded for, any major random change in the genetic code would have been lethal; hence it became "frozen".<sup id="cite_ref-pmid4887876_89-0" class="reference"><a href="#cite_note-pmid4887876-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup></li> <li>Stereochemical affinity: the genetic code is a result of a high affinity between each amino acid and its codon or anti-codon; the latter option implies that pre-tRNA molecules matched their corresponding amino acids by this affinity. Later during evolution, this matching was gradually replaced with matching by aminoacyl-tRNA synthetases.<sup id="cite_ref-pmid21779963_87-1" class="reference"><a href="#cite_note-pmid21779963-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid279919_90-0" class="reference"><a href="#cite_note-pmid279919-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19795157_91-0" class="reference"><a href="#cite_note-pmid19795157-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup></li> <li>Optimality: the genetic code continued to evolve after its initial creation, so that the current code maximizes some <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> function, usually some kind of error minimization.<sup id="cite_ref-pmid21779963_87-2" class="reference"><a href="#cite_note-pmid21779963-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10742043_88-1" class="reference"><a href="#cite_note-pmid10742043-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup></li></ul> <p>Hypotheses have addressed a variety of scenarios:<sup id="cite_ref-pmid10366854_93-0" class="reference"><a href="#cite_note-pmid10366854-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> </p> <ul><li>Chemical principles govern specific RNA interaction with amino acids. Experiments with <a href="/wiki/Aptamer" title="Aptamer">aptamers</a> showed that some amino acids have a selective chemical affinity for their codons.<sup id="cite_ref-pmid9751648_94-0" class="reference"><a href="#cite_note-pmid9751648-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> Experiments showed that of 8 amino acids tested, 6 show some RNA triplet-amino acid association.<sup id="cite_ref-isbn0-674-05075-4_81-1" class="reference"><a href="#cite_note-isbn0-674-05075-4-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19795157_91-1" class="reference"><a href="#cite_note-pmid19795157-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup></li> <li>Biosynthetic expansion. The genetic code grew from a simpler earlier code through a process of "biosynthetic expansion". Primordial life "discovered" new amino acids (for example, as by-products of <a href="/wiki/Metabolism" title="Metabolism">metabolism</a>) and later incorporated some of these into the machinery of genetic coding.<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> Although much circumstantial evidence has been found to suggest that fewer amino acid types were used in the past,<sup id="cite_ref-pmid12270892_96-0" class="reference"><a href="#cite_note-pmid12270892-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> precise and detailed hypotheses about which amino acids entered the code in what order are controversial.<sup id="cite_ref-pmid9115171_97-0" class="reference"><a href="#cite_note-pmid9115171-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11087835_98-0" class="reference"><a href="#cite_note-pmid11087835-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> However, several studies have suggested that Gly, Ala, Asp, Val, Ser, Pro, Glu, Leu, Thr may belong to a group of early-addition amino acids, whereas Cys, Met, Tyr, Trp, His, Phe may belong to a group of later-addition amino acids.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup></li> <li>Natural selection has led to codon assignments of the genetic code that minimize the effects of <a href="/wiki/Mutation" title="Mutation">mutations</a>.<sup id="cite_ref-pmid14604186_103-0" class="reference"><a href="#cite_note-pmid14604186-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> A recent hypothesis<sup id="cite_ref-pmid19479032_104-0" class="reference"><a href="#cite_note-pmid19479032-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> suggests that the triplet code was derived from codes that used longer than triplet codons (such as quadruplet codons). Longer than triplet decoding would increase codon redundancy and would be more error resistant. This feature could allow accurate decoding absent complex translational machinery such as the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>, such as before cells began making ribosomes.</li> <li>Information channels: <a href="/wiki/Information_theory" title="Information theory">Information-theoretic</a> approaches model the process of translating the genetic code into corresponding amino acids as an error-prone information channel.<sup id="cite_ref-pmid17826800_105-0" class="reference"><a href="#cite_note-pmid17826800-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> The inherent noise (that is, the error) in the channel poses the organism with a fundamental question: how can a genetic code be constructed to withstand noise<sup id="cite_ref-106" class="reference"><a href="#cite_note-106"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> while accurately and efficiently translating information? These <a href="/wiki/Rate-distortion_theory" class="mw-redirect" title="Rate-distortion theory">"rate-distortion"</a> models<sup id="cite_ref-pmid_18352335_107-0" class="reference"><a href="#cite_note-pmid_18352335-107"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> suggest that the genetic code originated as a result of the interplay of the three conflicting evolutionary forces: the needs for diverse amino acids,<sup id="cite_ref-pmid16838217_108-0" class="reference"><a href="#cite_note-pmid16838217-108"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> for error-tolerance<sup id="cite_ref-pmid14604186_103-1" class="reference"><a href="#cite_note-pmid14604186-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> and for minimal resource cost. The code emerges at a transition when the mapping of codons to amino acids becomes nonrandom. The code's emergence is governed by the <a href="/wiki/Topology" title="Topology">topology</a> defined by the probable errors and is related to the <a href="/wiki/Map_coloring_problem" class="mw-redirect" title="Map coloring problem">map coloring problem</a>.<sup id="cite_ref-pmid_20558115_109-0" class="reference"><a href="#cite_note-pmid_20558115-109"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup></li> <li>Game theory: Models based on <a href="/wiki/Signaling_game" title="Signaling game">signaling games</a> combine elements of game theory, natural selection and information channels. Such models have been used to suggest that the first polypeptides were likely short and had non-enzymatic function. Game theoretic models suggested that the organization of RNA strings into cells may have been necessary to prevent "deceptive" use of the genetic code, i.e. preventing the ancient equivalent of viruses from overwhelming the RNA world.<sup id="cite_ref-pmid23985735_110-0" class="reference"><a href="#cite_note-pmid23985735-110"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup></li> <li>Stop codons: Codons for translational stops are also an interesting aspect to the problem of the origin of the genetic code. As an example for addressing stop codon evolution, it has been suggested that the stop codons are such that they are most likely to terminate translation early in the case of a <a href="/wiki/Frame_shift" class="mw-redirect" title="Frame shift">frame shift</a> error.<sup id="cite_ref-111" class="reference"><a href="#cite_note-111"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> In contrast, some stereochemical molecular models explain the origin of stop codons as "unassignable".<sup id="cite_ref-pmid21779963_87-3" class="reference"><a href="#cite_note-pmid21779963-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup></li></ul> <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=Genetic_code&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/List_of_genetic_engineering_software" title="List of genetic engineering software">List of genetic engineering software</a></li> <li><a href="/wiki/Codon_tables" class="mw-redirect" title="Codon tables">Codon tables</a></li></ul> <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=Genetic_code&action=edit&section=16" title="Edit section: References"><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"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-pmid19131629-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid19131629_1-0">^</a></b></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 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The Royal Swedish Academy of Science. 1959<span class="reference-accessdate">. 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title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3785830">3785830</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/23985735">23985735</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+Royal+Society%2C+Interface&rft.atitle=What+can+information-asymmetric+games+tell+us+about+the+context+of+Crick%27s+%27frozen+accident%27%3F&rft.volume=10&rft.issue=88&rft.pages=20130614&rft.date=2013-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3785830%23id-name%3DPMC&rft_id=info%3Apmid%2F23985735&rft_id=info%3Adoi%2F10.1098%2Frsif.2013.0614&rft.aulast=Jee&rft.aufirst=J&rft.au=Sundstrom%2C+A&rft.au=Massey%2C+SE&rft.au=Mishra%2C+B&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3785830&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></span> </li> <li id="cite_note-111"><span class="mw-cite-backlink"><b><a href="#cite_ref-111">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFItzkovitzAlon2007" class="citation journal cs1">Itzkovitz S, Alon U (2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1832087">"The genetic code is nearly optimal for allowing additional information within protein-coding sequences"</a>. <i>Genome Research</i>. <b>17</b> (4): <span class="nowrap">405–</span>412. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgr.5987307">10.1101/gr.5987307</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/PMC1832087">1832087</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/17293451">17293451</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genome+Research&rft.atitle=The+genetic+code+is+nearly+optimal+for+allowing+additional+information+within+protein-coding+sequences&rft.volume=17&rft.issue=4&rft.pages=%3Cspan+class%3D%22nowrap%22%3E405-%3C%2Fspan%3E412&rft.date=2007&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1832087%23id-name%3DPMC&rft_id=info%3Apmid%2F17293451&rft_id=info%3Adoi%2F10.1101%2Fgr.5987307&rft.aulast=Itzkovitz&rft.aufirst=S&rft.au=Alon%2C+U&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1832087&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=17" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGriffithsMillerSuzukiLewontin1999" class="citation book cs1">Griffiths AJ, Miller JH, Suzuki DT, Lewontin RC, Gilbert WM (1999). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowTOC&rid=iga.TOC"><i>An Introduction to genetic analysis</i></a> (7th ed.). San Francisco: W.H. Freeman. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7167-3771-1" title="Special:BookSources/978-0-7167-3771-1"><bdi>978-0-7167-3771-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=An+Introduction+to+genetic+analysis&rft.place=San+Francisco&rft.edition=7th&rft.pub=W.H.+Freeman&rft.date=1999&rft.isbn=978-0-7167-3771-1&rft.aulast=Griffiths&rft.aufirst=Anthony+J.+F.&rft.au=Miller%2C+Jeffrey+H.&rft.au=Suzuki%2C+David+T.&rft.au=Lewontin%2C+Richard+C.&rft.au=Gilbert%2C+William+M.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2Fbv.fcgi%3Fcall%3Dbv.View..ShowTOC%26rid%3Diga.TOC&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAlbertsJohnsonLewisRaff2002" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowTOC&rid=mboc4.TOC&depth=2"><i>Molecular biology of the cell</i></a> (4th ed.). New York: Garland Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-3218-3" title="Special:BookSources/978-0-8153-3218-3"><bdi>978-0-8153-3218-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+biology+of+the+cell&rft.place=New+York&rft.edition=4th&rft.pub=Garland+Science&rft.date=2002&rft.isbn=978-0-8153-3218-3&rft.aulast=Alberts&rft.aufirst=Bruce&rft.au=Johnson%2C+Alexander&rft.au=Lewis%2C+Julian&rft.au=Raff%2C+Martin&rft.au=Roberts%2C+Keith&rft.au=Walter%2C+Peter&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2Fbv.fcgi%3Fcall%3Dbv.View..ShowTOC%26rid%3Dmboc4.TOC%26depth%3D2&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFLodishBerkZipurskyMatsudaira2000" class="citation book cs1">Lodish HF, Berk A, Zipursky SL, Matsudaira P, Baltimore D, Darnell JE (2000). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/molecularcellbi000lodi"><i>Molecular cell biology</i></a></span> (4th ed.). San Francisco: W.H. Freeman. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780716737063" title="Special:BookSources/9780716737063"><bdi>9780716737063</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+cell+biology&rft.place=San+Francisco&rft.edition=4th&rft.pub=W.H.+Freeman&rft.date=2000&rft.isbn=9780716737063&rft.aulast=Lodish&rft.aufirst=Harvey+F.&rft.au=Berk%2C+Arnold&rft.au=Zipursky%2C+S.+Lawrence&rft.au=Matsudaira%2C+Paul&rft.au=Baltimore%2C+David&rft.au=Darnell%2C+James+E.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmolecularcellbi000lodi&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFCaskeyLeder2014" class="citation journal cs1">Caskey CT, Leder P (April 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4000803">"The RNA code: nature's Rosetta Stone"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>111</b> (16): <span class="nowrap">5758–</span>9. <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/2014PNAS..111.5758C">2014PNAS..111.5758C</a>. <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/10.1073%2Fpnas.1404819111">10.1073/pnas.1404819111</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/PMC4000803">4000803</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/24756939">24756939</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=The+RNA+code%3A+nature%27s+Rosetta+Stone&rft.volume=111&rft.issue=16&rft.pages=%3Cspan+class%3D%22nowrap%22%3E5758-%3C%2Fspan%3E9&rft.date=2014-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4000803%23id-name%3DPMC&rft_id=info%3Apmid%2F24756939&rft_id=info%3Adoi%2F10.1073%2Fpnas.1404819111&rft_id=info%3Abibcode%2F2014PNAS..111.5758C&rft.aulast=Caskey&rft.aufirst=CT&rft.au=Leder%2C+P&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4000803&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+code" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_code&action=edit&section=18" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Commons-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/40px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/60px-Commons-logo.svg.png 1.5x" data-file-width="1024" data-file-height="1376" /></a></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Genetic_code" class="extiw" title="commons:Category:Genetic code">Genetic code</a></span>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/taxonomyhome.html/index.cgi?chapter=cgencodes">The Genetic Codes: Genetic Code Tables</a></li> <li>The <a rel="nofollow" class="external text" href="http://www.kazusa.or.jp/codon/">Codon Usage Database</a> — Codon frequency tables for many organisms</li> <li><a rel="nofollow" class="external text" href="http://history.nih.gov/exhibits/nirenberg/">History of deciphering the genetic code</a></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 dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul 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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" /><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 ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini 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template">e</abbr></a></li></ul></div><div id="Genetics421" style="font-size:114%;margin:0 4em"><a href="/wiki/Genetics" title="Genetics">Genetics</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Timeline_of_the_history_of_genetics" title="Timeline of the history of genetics">Timeline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li> <li><a href="/wiki/Glossary_of_genetics" class="mw-redirect" title="Glossary of genetics">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key components</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/Chromosome" title="Chromosome">Chromosome</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotide</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</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/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a> of</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/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a></li> <li><a href="/wiki/Genetic_history_of_indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of indigenous peoples of the Americas">the Americas</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</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/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a class="mw-selflink selflink">Genetic code</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Genetic diversity</a></li> <li><a href="/wiki/Genetic_monitoring" title="Genetic monitoring">Genetic monitoring</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/He_Jiankui_genome_editing_incident" class="mw-redirect" title="He Jiankui genome editing incident">He Jiankui genome editing incident</a></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</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/Plant_genetics" title="Plant genetics">Plant genetics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></li> <li><a href="/wiki/List_of_genetics_research_organizations" title="List of genetics research organizations">List of genetics research organizations</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:Genetics" title="Category:Genetics">Category</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="Gene_expression217" 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:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression217" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction<br />to genetics</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 class="mw-selflink selflink">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a href="/wiki/DNA" title="DNA">DNA</a> → <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a> → <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</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:8.0em;font-weight: normal;">Types</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/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</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/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</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/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA / hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Translation_(biology)" title="Translation (biology)">Translation</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:8.0em;font-weight: normal;">Types</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/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</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/Epigenetics" title="Epigenetics">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</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/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</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" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /></div><div role="navigation" class="navbox" aria-labelledby="Encoded_(proteinogenic)_amino_acids563" 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="3"><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:Proteinogenic_amino_acids" title="Template:Proteinogenic amino acids"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Proteinogenic_amino_acids" title="Template talk:Proteinogenic amino acids"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Proteinogenic_amino_acids" title="Special:EditPage/Template:Proteinogenic amino acids"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Encoded_(proteinogenic)_amino_acids563" style="font-size:114%;margin:0 4em"><a href="/wiki/Proteinogenic_amino_acid" title="Proteinogenic amino acid">Encoded (proteinogenic) amino acids</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">General topics</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/Protein" title="Protein">Protein</a></li> <li><a href="/wiki/Peptide" title="Peptide">Peptide</a></li> <li><a class="mw-selflink selflink">Genetic code</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="2" style="width:1px;padding:0 0 0 2px"><div><figure typeof="mw:File/Thumb"><a href="/wiki/File:L-amino_acid_any.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/L-amino_acid_any.png/120px-L-amino_acid_any.png" decoding="async" width="100" height="88" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/L-amino_acid_any.png/250px-L-amino_acid_any.png 1.5x" data-file-width="728" data-file-height="640" /></a><figcaption>Unspecified L-amino acid</figcaption></figure></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By properties</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%"><a href="/wiki/Aliphatic_compound" title="Aliphatic compound">Aliphatic</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/Branched-chain_amino_acid" title="Branched-chain amino acid">Branched-chain amino acids</a> (<a href="/wiki/Valine" title="Valine">Valine</a></li> <li><a href="/wiki/Isoleucine" title="Isoleucine">Isoleucine</a></li> <li><a href="/wiki/Leucine" title="Leucine">Leucine</a>)</li> <li><a href="/wiki/Methionine" title="Methionine">Methionine</a></li> <li><a href="/wiki/Alanine" title="Alanine">Alanine</a></li> <li><a href="/wiki/Proline" title="Proline">Proline</a></li> <li><a href="/wiki/Glycine" title="Glycine">Glycine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Aromatic_amino_acid" title="Aromatic amino acid">Aromatic</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/Histidine" title="Histidine">Histidine</a></li> <li><a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a></li> <li><a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</a></li> <li><a href="/wiki/Phenylalanine" title="Phenylalanine">Phenylalanine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Chemical_polarity" title="Chemical polarity">Polar</a>, uncharged</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/Asparagine" title="Asparagine">Asparagine</a></li> <li><a href="/wiki/Glutamine" title="Glutamine">Glutamine</a></li> <li><a href="/wiki/Serine" title="Serine">Serine</a></li> <li><a href="/wiki/Threonine" title="Threonine">Threonine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Positive charge (p<i>K</i><sub>a</sub>)</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/Lysine" title="Lysine">Lysine</a> (≈10.8)</li> <li><a href="/wiki/Arginine" title="Arginine">Arginine</a> (≈12.5)</li> <li><a href="/wiki/Histidine" title="Histidine">Histidine</a> (≈6.1)</li> <li><a href="/wiki/Pyrrolysine" title="Pyrrolysine">Pyrrolysine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Negative charge (p<i>K</i><sub>a</sub>)</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/Aspartic_acid" title="Aspartic acid">Aspartic acid</a> (≈3.9)</li> <li><a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> (≈4.1)</li> <li><a href="/wiki/Selenocysteine" title="Selenocysteine">Selenocysteine</a> (≈5.4)</li> <li><a href="/wiki/Cysteine" title="Cysteine">Cysteine</a> (≈8.3)</li> <li><a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a> (≈10.1)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div><div class="hlist" style="font-size:88%;"> <ul><li><b><a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">Amino acids</a> types</b>: <a href="/wiki/Template:Encoded_amino_acids" class="mw-redirect" title="Template:Encoded amino acids">Encoded (proteins)</a></li> <li><a href="/wiki/Essential_amino_acids" class="mw-redirect" title="Essential amino acids">Essential</a></li> <li><a href="/wiki/Template:Non-proteinogenic_amino_acids" title="Template:Non-proteinogenic amino acids">Non-proteinogenic</a></li> <li><a href="/wiki/Ketogenic_amino_acids" class="mw-redirect" title="Ketogenic amino acids">Ketogenic</a></li> <li><a href="/wiki/Glucogenic_amino_acids" class="mw-redirect" title="Glucogenic amino acids">Glucogenic</a></li> <li><a href="/wiki/Secondary_amino_acid" title="Secondary amino acid">Secondary amino</a></li> <li><a href="/wiki/Imino_acid" title="Imino acid">Imino acids</a></li> <li><a href="/wiki/D-amino_acids" class="mw-redirect" title="D-amino acids">D-amino acids</a></li> <li><a href="/wiki/Dehydroamino_acid" title="Dehydroamino acid">Dehydroamino acids</a></li></ul> </div></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="Biochemistry_topics442" 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:Biochemistry_topics" title="Template:Biochemistry topics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biochemistry_topics" title="Template talk:Biochemistry topics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Biochemistry_topics" title="Special:EditPage/Template:Biochemistry topics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Biochemistry_topics442" style="font-size:114%;margin:0 4em"><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a> topics</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Science fields</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/List_of_life_sciences" title="List of life sciences">List of life sciences</a></li> <li><a href="/wiki/Chemistry" title="Chemistry">Chemistry</a></li> <li><a href="/wiki/Biology" title="Biology">Biology</a></li> <li><i>Chemistry fields:</i></li> <li><a href="/wiki/Analytical_chemistry" title="Analytical chemistry">Analytical chemistry</a></li> <li><a href="/wiki/Inorganic_chemistry" title="Inorganic chemistry">Inorganic chemistry</a></li> <li><a href="/wiki/Medicinal_chemistry" title="Medicinal chemistry">Medicinal chemistry</a></li> <li><a href="/wiki/Organic_chemistry" title="Organic chemistry">Organic chemistry</a></li> <li><a href="/wiki/Physical_chemistry" title="Physical chemistry">Physical chemistry</a></li> <li><a href="/wiki/Theoretical_chemistry" title="Theoretical chemistry">Theoretical chemistry</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a><br /> fields</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/Molecular_biology" title="Molecular biology">Molecular biology</a></li> <li><a href="/wiki/Cell_biology" title="Cell biology">Cell biology</a></li> <li><a href="/wiki/Chemical_biology" title="Chemical biology">Chemical biology</a> <ul><li><a href="/wiki/Bioorthogonal_chemistry" title="Bioorthogonal chemistry">Bioorthogonal chemistry</a></li></ul></li> <li><a href="/wiki/Medicinal_chemistry" title="Medicinal chemistry">Medicinal chemistry</a> <ul><li><a href="/wiki/Pharmacology" title="Pharmacology">Pharmacology</a></li></ul></li> <li><a href="/wiki/Clinical_chemistry" title="Clinical chemistry">Clinical chemistry</a></li> <li><a href="/wiki/Neurochemistry" title="Neurochemistry">Neurochemistry</a></li> <li><a href="/wiki/Bioorganic_chemistry" title="Bioorganic chemistry">Bioorganic chemistry</a></li> <li><a href="/wiki/Bioorganometallic_chemistry" title="Bioorganometallic chemistry">Bioorganometallic chemistry</a></li> <li><a href="/wiki/Bioinorganic_chemistry" title="Bioinorganic chemistry">Bioinorganic chemistry</a></li> <li><a href="/wiki/Biophysical_chemistry" title="Biophysical chemistry">Biophysical chemistry</a></li> <li><a href="/wiki/Bacteriology" title="Bacteriology">Bacteriology</a></li> <li><a href="/wiki/Parasitology" title="Parasitology">parasitology</a></li> <li><a href="/wiki/Virology" title="Virology">virology</a></li> <li><a href="/wiki/Immunology" title="Immunology">immunology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General <br /> chemistry</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/Chemical_reactions" class="mw-redirect" title="Chemical reactions">Chemical reactions</a></li> <li><a href="/wiki/Chemical_synthesis" title="Chemical synthesis">Chemical synthesis</a></li> <li><a href="/wiki/Molecule" title="Molecule">Molecule</a></li> <li><a href="/wiki/Organic_reactions" class="mw-redirect" title="Organic reactions">Organic reactions</a></li> <li><a href="/wiki/Catalyst" class="mw-redirect" title="Catalyst">Catalyst</a></li> <li><a href="/wiki/Reagent" title="Reagent">Reagent</a></li> <li><a href="/wiki/Reaction_mechanism" title="Reaction mechanism">Reaction mechanism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Concepts</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/Structural_biology" title="Structural biology">Structural biology</a></li> <li><a href="/wiki/Enzymology" class="mw-redirect" title="Enzymology">Enzymology</a></li> <li><a href="/wiki/Metabolism" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Biomolecule" title="Biomolecule">Biomolecule</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cells</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><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_theory" title="Cell theory">Cell theory</a></li> <li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></li> <li><a href="/wiki/Peptide" title="Peptide">Peptide</a></li> <li><a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">Amino acids</a></li> <li><a href="/wiki/Protein" title="Protein">Protein</a></li> <li><a href="/wiki/Peptide" title="Peptide">Peptide</a></li> <li><a href="/wiki/Hormone" title="Hormone">Hormone</a></li> <li><a href="/wiki/Enzyme" title="Enzyme">Enzyme</a></li> <li><a href="/wiki/Metabolic_pathway" title="Metabolic pathway">Metabolic pathway</a></li></ul> <ul><li><i><a href="/wiki/Genetics" title="Genetics">Genetics</a>:</i></li> <li><a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">Chromosomes</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a></li> <li><a class="mw-selflink selflink">Genetic code</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Biomolecule" title="Biomolecule">Biomolecule</a> <br /> families</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><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><u><a href="/wiki/Carbohydrates" class="mw-redirect" title="Carbohydrates">Carbohydrates</a>:</u></i></li> <li><a href="/wiki/Alcohols" class="mw-redirect" title="Alcohols">Alcohols</a></li> <li><a href="/wiki/Glycoproteins" class="mw-redirect" title="Glycoproteins">Glycoproteins</a></li> <li><a href="/wiki/Glycosides" class="mw-redirect" title="Glycosides">Glycosides</a></li> <li><i><u><a href="/wiki/Lipids" class="mw-redirect" title="Lipids">Lipids</a>:</u></i></li> <li><a href="/wiki/Eicosanoids" class="mw-redirect" title="Eicosanoids">Eicosanoids</a></li> <li><a href="/wiki/Fatty_acids" class="mw-redirect" title="Fatty acids">Fatty acids</a> <ul><li><a href="/wiki/Fatty-acid_metabolism" class="mw-redirect" title="Fatty-acid metabolism">Fatty-acid metabolism</a></li></ul></li> <li><a href="/wiki/Glycerides" class="mw-redirect" title="Glycerides">Glycerides</a></li> <li><a href="/wiki/Phospholipids" class="mw-redirect" title="Phospholipids">Phospholipids</a></li> <li><a href="/wiki/Sphingolipids" class="mw-redirect" title="Sphingolipids">Sphingolipids</a></li> <li><a href="/wiki/Cholesterol" title="Cholesterol">Cholesterol</a></li> <li><a href="/wiki/Steroids" class="mw-redirect" title="Steroids">Steroids</a></li> <li><u><i><a href="/wiki/Nucleic_acids" class="mw-redirect" title="Nucleic acids">Nucleic acids</a>:</i></u></li> <li><a href="/wiki/Nucleobases" class="mw-redirect" title="Nucleobases">Nucleobases</a></li> <li><a href="/wiki/Nucleosides" class="mw-redirect" title="Nucleosides">Nucleosides</a></li> <li><a href="/wiki/Nucleotides" class="mw-redirect" title="Nucleotides">Nucleotides</a> <ul><li><a href="/wiki/Nucleotide_metabolism" class="mw-redirect" title="Nucleotide metabolism">Nucleotide metabolism</a></li></ul></li> <li><i><u><a href="/wiki/Proteins" class="mw-redirect" title="Proteins">Proteins</a>:</u></i></li> <li><a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">Amino acids</a> <ul><li><a href="/wiki/Amino_acid_metabolism" class="mw-redirect" title="Amino acid metabolism">Amino acid metabolism</a></li></ul></li> <li><i><u>Other:</u></i></li> <li><a href="/wiki/Tetrapyrroles" class="mw-redirect" title="Tetrapyrroles">Tetrapyrroles</a></li> <li><a href="/wiki/Heme" title="Heme">Heme</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Chemical_synthesis" title="Chemical synthesis">Chemical <br /> synthesis</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/Artificial_gene_synthesis" title="Artificial gene synthesis">Artificial gene synthesis</a></li> <li><a href="/wiki/Biomimetic_synthesis" title="Biomimetic synthesis">Biomimetic synthesis</a></li> <li><a href="/wiki/Bioretrosynthesis" title="Bioretrosynthesis">Bioretrosynthesis</a></li> <li><a href="/wiki/Biosynthesis" title="Biosynthesis">Biosynthesis</a></li> <li><a href="/wiki/Chemosynthesis_(nanotechnology)" title="Chemosynthesis (nanotechnology)">Chemosynthesis</a></li> <li><a href="/wiki/Convergent_synthesis" title="Convergent synthesis">Convergent synthesis</a></li> <li><a href="/wiki/Custom_peptide_synthesis" title="Custom peptide synthesis">Custom peptide synthesis</a></li> <li><a href="/wiki/Direct_process" title="Direct process">Direct process</a></li> <li><a href="/wiki/Divergent_synthesis" title="Divergent synthesis">Divergent synthesis</a></li> <li><a href="/wiki/Electrosynthesis" title="Electrosynthesis">Electrosynthesis</a></li> <li><a href="/wiki/Enantioselective_synthesis" title="Enantioselective synthesis">Enantioselective synthesis</a></li> <li><a href="/wiki/Fully_automated_synthesis" class="mw-redirect" title="Fully automated synthesis">Fully automated synthesis</a></li> <li><a href="/wiki/Hydrothermal_synthesis" title="Hydrothermal synthesis">Hydrothermal synthesis</a></li> <li><a href="/wiki/Laser_ablation_synthesis_in_solution" title="Laser ablation synthesis in solution">LASiS</a></li> <li><a href="/wiki/Mechanosynthesis" title="Mechanosynthesis">Mechanosynthesis</a></li> <li><a href="/wiki/One-pot_synthesis" title="One-pot synthesis">One-pot synthesis</a></li> <li><a href="/wiki/Organic_synthesis" title="Organic synthesis">Organic synthesis</a></li> <li><a href="/wiki/Peptide_synthesis" title="Peptide synthesis">Peptide synthesis</a></li> <li><a href="/wiki/Radiosynthesis" title="Radiosynthesis">Radiosynthesis</a></li> <li><a href="/wiki/Retrosynthetic_analysis" title="Retrosynthetic analysis">Retrosynthesis</a></li> <li><a href="/wiki/Semisynthesis" title="Semisynthesis">Semisynthesis</a></li> <li><a href="/wiki/Solid-phase_synthesis" title="Solid-phase synthesis">Solid-phase synthesis</a></li> <li><a href="/wiki/Solvothermal_synthesis" title="Solvothermal synthesis">Solvothermal synthesis</a></li> <li><a href="/wiki/Total_synthesis" title="Total synthesis">Total synthesis</a></li> <li><a href="/wiki/Volume_combustion_synthesis" title="Volume combustion synthesis">Volume combustion synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Templates</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span class="nowrap">{{</span><a href="/wiki/Template:Branches_of_chemistry" title="Template:Branches of chemistry">Branches of chemistry</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Cellular_structures" title="Template:Cellular structures">Cellular structures</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Organic_reactions" title="Template:Organic reactions">Organic reactions</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Chemical_synthesis" title="Template:Chemical synthesis">Chemical synthesis</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Amino_acids" class="mw-redirect" title="Template:Amino acids">Amino acids</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Chemical_bonds" title="Template:Chemical bonds">Chemical bonds</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Protein_primary_structure" title="Template:Protein primary structure">Protein primary structure</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Amino_acid_metabolism_enzymes" title="Template:Amino acid metabolism enzymes">Amino acid metabolism enzymes</a><span class="nowrap">}}</span></li></ul> <ul><li><span class="nowrap">{{</span><a href="/wiki/Template:Life_on_Earth" title="Template:Life on Earth">Life on Earth</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Botany" title="Template:Botany">Botany</a><span class="nowrap">}}</span></li> <li><span class="nowrap">{{</span><a href="/wiki/Template:Zoology" title="Template:Zoology">Zoology</a><span class="nowrap">}}</span></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><i>Occupations</i></li> <li><a href="/wiki/Biologist" title="Biologist">Biologist</a></li> <li><a href="/wiki/Chemist" title="Chemist">Chemist</a></li> <li><a href="/wiki/Bacteriologist" title="Bacteriologist">Bacteriologist</a></li> <li><a href="/wiki/Microbiologist" title="Microbiologist">Microbiologist</a></li> <li><a href="/wiki/Microscopy" title="Microscopy">Microscopy</a></li> <li><i>Concepts:</i></li> <li><a href="/wiki/Scientific_method" title="Scientific method">Scientific method</a></li> <li><a href="/wiki/Periodic_table" title="Periodic table">Periodic table</a></li> <li><a href="/wiki/Experiments" class="mw-redirect" title="Experiments">Experiments</a></li> <li><i>Notable scientists:</i></li> <li><a href="/wiki/Robert_Hooke" title="Robert Hooke">Robert Hooke</a></li> <li><a href="/wiki/Antonie_van_Leeuwenhoek" title="Antonie van Leeuwenhoek">Antonie van Leeuwenhoek</a></li> <li><a href="/wiki/Francesco_Redi" title="Francesco Redi">Francesco Redi</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" 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