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<span>Overview</span> </div> </a> <ul id="toc-Overview-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_a_unit_of_transcription" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#As_a_unit_of_transcription"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>As a unit of transcription</span> </div> </a> <ul id="toc-As_a_unit_of_transcription-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Operons_versus_clustering_of_prokaryotic_genes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Operons_versus_clustering_of_prokaryotic_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Operons versus clustering of prokaryotic genes</span> </div> </a> <ul id="toc-Operons_versus_clustering_of_prokaryotic_genes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-General_structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#General_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>General structure</span> </div> </a> <ul id="toc-General_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Regulation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Regulation</span> </div> </a> <ul id="toc-Regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-The_lac_operon" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#The_lac_operon"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>The <i>lac</i> operon</span> </div> </a> <ul id="toc-The_lac_operon-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-The_trp_operon" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#The_trp_operon"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>The <i>trp</i> operon</span> </div> </a> <ul id="toc-The_trp_operon-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Predicting_the_number_and_organization_of_operons" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Predicting_the_number_and_organization_of_operons"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Predicting the number and organization of operons</span> </div> </a> <ul id="toc-Predicting_the_number_and_organization_of_operons-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">10</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">11</span> <span>References</span> </div> </a> <ul id="toc-References-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">12</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" > <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">Operon</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 44 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-44" 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">44 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/%D9%85%D8%B4%D8%BA%D9%84_(%D8%A3%D8%AD%D9%8A%D8%A7%D8%A1)" title="مشغل (أحياء) – Arabic" lang="ar" hreflang="ar" data-title="مشغل (أحياء)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Bulgarian" lang="bg" hreflang="bg" data-title="Оперон" 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/Operon" title="Operon – Bosnian" lang="bs" hreflang="bs" data-title="Operon" 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/Oper%C3%B3" title="Operó – Catalan" lang="ca" hreflang="ca" data-title="Operó" 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/Operon" title="Operon – Czech" lang="cs" hreflang="cs" data-title="Operon" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Operon" title="Operon – German" lang="de" hreflang="de" data-title="Operon" 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/Operon" title="Operon – Estonian" lang="et" hreflang="et" data-title="Operon" 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%9F%CF%80%CE%B5%CF%81%CF%8C%CE%BD%CE%B9%CE%BF" title="Οπερόνιο – Greek" lang="el" hreflang="el" data-title="Οπερόνιο" 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/Oper%C3%B3n" title="Operón – Spanish" lang="es" hreflang="es" data-title="Operón" 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/Operono" title="Operono – Esperanto" lang="eo" hreflang="eo" data-title="Operono" 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/Operoi" title="Operoi – Basque" lang="eu" hreflang="eu" data-title="Operoi" 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%A7%D9%BE%D8%B1%D9%88%D9%86" title="اپرون – Persian" lang="fa" hreflang="fa" data-title="اپرون" 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/Op%C3%A9ron" title="Opéron – French" lang="fr" hreflang="fr" data-title="Opéron" 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/Obr%C3%B3n" title="Obrón – Irish" lang="ga" hreflang="ga" data-title="Obrón" 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/Oper%C3%B3n" title="Operón – Galician" lang="gl" hreflang="gl" data-title="Operón" 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%98%A4%ED%8E%98%EB%A1%A0" title="오페론 – Korean" lang="ko" hreflang="ko" data-title="오페론" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%95%D5%BA%D5%A5%D6%80%D5%B8%D5%B6" title="Օպերոն – Armenian" lang="hy" hreflang="hy" data-title="Օպերոն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Operon" title="Operon – Croatian" lang="hr" hreflang="hr" data-title="Operon" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Operon" title="Operon – Indonesian" lang="id" hreflang="id" data-title="Operon" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Operone" title="Operone – Italian" lang="it" hreflang="it" data-title="Operone" 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%90%D7%95%D7%A4%D7%A8%D7%95%D7%9F" title="אופרון – Hebrew" lang="he" hreflang="he" data-title="אופרון" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%93%E0%B2%AA%E0%B2%B0%E0%B2%BE%E0%B2%A8%E0%B3%8D" title="ಓಪರಾನ್ – Kannada" lang="kn" hreflang="kn" data-title="ಓಪರಾನ್" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Kazakh" lang="kk" hreflang="kk" data-title="Оперон" 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/Operon" title="Operon – Kurdish" lang="ku" hreflang="ku" data-title="Operon" 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%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Kyrgyz" lang="ky" hreflang="ky" data-title="Оперон" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Operon-modell" title="Operon-modell – Hungarian" lang="hu" hreflang="hu" data-title="Operon-modell" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Operon" title="Operon – Dutch" lang="nl" hreflang="nl" data-title="Operon" 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/%E3%82%AA%E3%83%9A%E3%83%AD%E3%83%B3" title="オペロン – Japanese" lang="ja" hreflang="ja" data-title="オペロン" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Operon" title="Operon – Occitan" lang="oc" hreflang="oc" data-title="Operon" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Operon_(biologia)" title="Operon (biologia) – Polish" lang="pl" hreflang="pl" data-title="Operon (biologia)" 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/Oper%C3%A3o" title="Operão – Portuguese" lang="pt" hreflang="pt" data-title="Operão" 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/Operon" title="Operon – Romanian" lang="ro" hreflang="ro" data-title="Operon" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Russian" lang="ru" hreflang="ru" data-title="Оперон" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Oper%C3%B3n" title="Operón – Slovak" lang="sk" hreflang="sk" data-title="Operón" 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/Operon" title="Operon – Slovenian" lang="sl" hreflang="sl" data-title="Operon" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Serbian" lang="sr" hreflang="sr" data-title="Оперон" 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/Operon" title="Operon – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Operon" 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/Operoni" title="Operoni – Finnish" lang="fi" hreflang="fi" data-title="Operoni" 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/Operon" title="Operon – Swedish" lang="sv" hreflang="sv" data-title="Operon" 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%AE%A3%E0%AF%81%E0%AE%A4%E0%AF%8D%E0%AE%A4%E0%AF%8A%E0%AE%95%E0%AF%81%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AF%81" title="மரபணுத்தொகுப்பு – Tamil" lang="ta" hreflang="ta" data-title="மரபணுத்தொகுப்பு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Operon" title="Operon – Turkish" lang="tr" hreflang="tr" data-title="Operon" 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%9E%D0%BF%D0%B5%D1%80%D0%BE%D0%BD" title="Оперон – Ukrainian" lang="uk" hreflang="uk" data-title="Оперон" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Operon" title="Operon – Vietnamese" lang="vi" hreflang="vi" data-title="Operon" 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-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E6%93%8D%E7%B8%B1%E5%AD%90" title="操縱子 – Chinese" lang="zh" hreflang="zh" data-title="操縱子" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q139677#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-associated-pages" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-nstab-main" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Operon" title="View the content page [c]" accesskey="c"><span>Article</span></a></li><li id="ca-talk" class="vector-tab-noicon mw-list-item"><a href="/wiki/Talk:Operon" rel="discussion" title="Discuss improvements to the content page [t]" 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class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Group of open reading frames under the same regulation</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Opteron" title="Opteron">Opteron</a> or <a href="/wiki/Oberon" title="Oberon">Oberon</a>.</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Operon_1.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Operon_1.png/350px-Operon_1.png" decoding="async" width="350" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/a/a4/Operon_1.png 1.5x" data-file-width="490" data-file-height="59" /></a><figcaption>A typical operon</figcaption></figure> <p>In <a href="/wiki/Genetics" title="Genetics">genetics</a>, an <b>operon</b> is a functioning unit of <a href="/wiki/DNA" title="DNA">DNA</a> containing a cluster of <a href="/wiki/Gene" title="Gene">genes</a> under the control of a single <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a>.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The genes are <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcribed</a> together into an <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a> strand and either <a href="/wiki/Translation_(biology)" title="Translation (biology)">translated</a> together in the cytoplasm, or undergo <a href="/wiki/RNA_splicing" title="RNA splicing">splicing</a> to create <a href="/wiki/Cistron" title="Cistron">monocistronic</a> mRNAs that are translated separately, i.e. several strands of mRNA that each encode a single gene product. The result of this is that the genes contained in the operon are either <a href="/wiki/Gene_expression" title="Gene expression">expressed</a> together or not at all. Several genes must be <i>co-transcribed</i> to define an operon.<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> </p><p>Originally, operons were thought to exist solely in <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> (which includes <a href="/wiki/Organelle" title="Organelle">organelles</a> like <a href="/wiki/Plastid" title="Plastid">plastids</a> that are derived from <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>), but their discovery in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> was shown in the early 1990s, and are considered to be rare.<sup id="cite_ref-Kominek_3-0" class="reference"><a href="#cite_note-Kominek-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Operons_in_eukaryotes_6-0" class="reference"><a href="#cite_note-Operons_in_eukaryotes-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In general, expression of prokaryotic operons leads to the generation of <a href="/wiki/Cistron" title="Cistron">polycistronic</a> mRNAs, while eukaryotic operons lead to monocistronic mRNAs. </p><p>Operons are also found in viruses such as <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophages</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Liu_2004_8-0" class="reference"><a href="#cite_note-Liu_2004-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/T7_phage" title="T7 phage">T7 phages</a> have two operons. The first operon codes for various products, including a special <a href="/wiki/T7_RNA_polymerase" title="T7 RNA polymerase">T7 RNA polymerase</a> which can bind to and transcribe the second operon. The second operon includes a <a href="/wiki/Lysis" title="Lysis">lysis</a> gene meant to cause the host cell to burst.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <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=Operon&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term "operon" was first proposed in a short paper in the Proceedings of the <a href="/wiki/French_Academy_of_Science" class="mw-redirect" title="French Academy of Science">French Academy of Science</a> in 1960.<sup id="cite_ref-Jacob1960_10-0" class="reference"><a href="#cite_note-Jacob1960-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> From this paper, the so-called general theory of the operon was developed. This theory suggested that in all cases, genes within an operon are negatively controlled by a <a href="/wiki/Repressor" title="Repressor">repressor</a> acting at a single <a href="/wiki/Operator_(biology)" class="mw-redirect" title="Operator (biology)">operator</a> located before the first gene. Later, it was discovered that genes could be positively regulated and also regulated at steps that follow transcription initiation. Therefore, it is not possible to talk of a general regulatory mechanism, because different operons have different mechanisms. Today, the operon is simply defined as a cluster of genes transcribed into a single mRNA molecule. Nevertheless, the development of the concept is considered a landmark event in the history of molecular biology. The first operon to be described was the <a href="/wiki/Lac_operon" title="Lac operon"><i>lac</i> operon</a> in <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a></i>.<sup id="cite_ref-Jacob1960_10-1" class="reference"><a href="#cite_note-Jacob1960-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The 1965 <a href="/wiki/Nobel_Prize_in_Physiology_and_Medicine" class="mw-redirect" title="Nobel Prize in Physiology and Medicine">Nobel Prize in Physiology and Medicine</a> was awarded to <a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a>, <a href="/wiki/Andr%C3%A9_Michel_Lwoff" title="André Michel Lwoff">André Michel Lwoff</a> and <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> for their discoveries concerning the operon and virus synthesis. </p> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=2" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table role="presentation" cellpadding="0" style="border-spacing:0; clear:right; float:right;"> <tbody><tr> <td><div class="thumb tright"> <div class="thumbinner" style="width:602px;"> <div style="position:relative; width:600px; height:300px; overflow:hidden; border:solid #ccc 1px; background-color:white;"> <div style="left:0px; top:0px; width:600px; position:absolute"> <span typeof="mw:File"><a href="/wiki/File:Gene_structure_prokaryote_2_annotated.svg" title="File:Gene structure prokaryote 2 annotated.svg"><img alt="Prokaryote gene structure diagram" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Gene_structure_prokaryote_2_unannotated.svg/600px-Gene_structure_prokaryote_2_unannotated.svg.png" decoding="async" width="600" height="295" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Gene_structure_prokaryote_2_unannotated.svg/900px-Gene_structure_prokaryote_2_unannotated.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Gene_structure_prokaryote_2_unannotated.svg/1200px-Gene_structure_prokaryote_2_unannotated.svg.png 2x" data-file-width="1190" data-file-height="586" /></a></span> </div> <div style="text-align:left; font-size:11px; line-height:110%"> <div style="background-color:transparent; color:black"><div id="annotation_310x2" style="position:absolute; left:310px; top:2px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a class="mw-selflink selflink">Polycistronic operon</a> </span></div> <div id="annotation_115x37" style="position:absolute; left:115px; top:37px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_475x37" style="position:absolute; left:475px; top:37px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_71x66" style="position:absolute; left:71px; top:66px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_535x66" style="position:absolute; left:535px; top:66px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_535x77" style="position:absolute; left:535px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_71x77" style="position:absolute; left:71px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_139x77" style="position:absolute; left:139px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Operator_(genetics)" class="mw-redirect" title="Operator (genetics)">Operator</a> </span></div> <div id="annotation_187x77" style="position:absolute; left:187px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> </span></div> <div id="annotation_236x77" style="position:absolute; left:236px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/5%27UTR" class="mw-redirect" title="5'UTR">5'UTR </a> </span></div> <div id="annotation_301x77" style="position:absolute; left:301px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Open_reading_frame" title="Open reading frame">ORF </a> </span></div> <div id="annotation_431x77" style="position:absolute; left:431px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Open_reading_frame" title="Open reading frame">ORF </a> </span></div> <div id="annotation_375x77" style="position:absolute; left:375px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Untranslated_region" title="Untranslated region">UTR </a> </span></div> <div id="annotation_475x77" style="position:absolute; left:475px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/3%27UTR" class="mw-redirect" title="3'UTR">3'UTR </a> </span></div> <div id="annotation_268x103" style="position:absolute; left:268px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Start_codon" title="Start codon">Start </a> </span></div> <div id="annotation_410x103" style="position:absolute; left:410px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Start_codon" title="Start codon">Start </a> </span></div> <div id="annotation_344x103" style="position:absolute; left:344px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Stop_codon" title="Stop codon">Stop </a> </span></div> <div id="annotation_450x103" style="position:absolute; left:450px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Stop_codon" title="Stop codon">Stop </a> </span></div> <div id="annotation_483x103" style="position:absolute; left:483px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">Terminator</a> </span></div> <div id="annotation_51x155" style="position:absolute; left:51px; top:155px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription</a> </span></div> <div id="annotation_18x124" style="position:absolute; left:18px; top:124px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/DNA" title="DNA">DNA </a> </span></div> <div id="annotation_239x170" style="position:absolute; left:239px; top:170px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Ribosome_binding_site" class="mw-redirect" title="Ribosome binding site">RBS </a> </span></div> <div id="annotation_382x170" style="position:absolute; left:382px; top:170px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Ribosome_binding_site" class="mw-redirect" title="Ribosome binding site">RBS </a> </span></div> <div id="annotation_264x156" style="position:absolute; left:264px; top:156px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">Protein coding region </a> </span></div> <div id="annotation_408x156" style="position:absolute; left:408px; top:156px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">Protein coding region </a> </span></div> <div id="annotation_14x195" style="position:absolute; left:14px; top:195px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA </a> </span></div> <div id="annotation_51x219" style="position:absolute; left:51px; top:219px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">Translation</a> </span></div> <div id="annotation_14x256" style="position:absolute; left:14px; top:256px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein" title="Protein">Protein </a> </span></div></div> </div> </div> <div class="thumbcaption" style="clear:left"><div style="float:left;margin-right:0.5em"><span typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/18px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/27px-Interactive_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/36px-Interactive_icon.svg.png 2x" data-file-width="133" data-file-height="200" /></span></span></div>The structure of a <a href="/wiki/Prokaryotic" class="mw-redirect" title="Prokaryotic">prokaryotic</a> <a class="mw-selflink selflink">operon</a> of protein-coding <a href="/wiki/Gene" title="Gene">genes</a>. <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence</a> controls when expression occurs for the multiple <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">protein coding regions</a> (red). <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a>, <a href="/wiki/Operator_(biology)" class="mw-redirect" title="Operator (biology)">operator</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions (yellow) regulate the <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of the gene into an mRNA. The mRNA <a href="/wiki/Untranslated_region" title="Untranslated region">untranslated regions</a> (blue) regulate <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> into the final protein products.<sup id="cite_ref-ShafeeLowe2017_11-0" class="reference"><a href="#cite_note-ShafeeLowe2017-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup></div> </div> </div> </td></tr></tbody></table> <p>Operons occur primarily in <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> but also rarely in some <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, including <a href="/wiki/Nematode" title="Nematode">nematodes</a> such as <a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans"><i>C. elegans</i></a> and the fruit fly, <a href="/wiki/Drosophila" title="Drosophila"><i>Drosophila melanogaster</i></a>.<sup id="cite_ref-Kominek_3-1" class="reference"><a href="#cite_note-Kominek-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">rRNA</a> genes often exist in operons that have been found in a range of eukaryotes including <a href="/wiki/Chordate" title="Chordate">chordates</a>. An operon is made up of several <a href="/wiki/Structural_gene" title="Structural gene">structural genes</a> arranged under a common <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a> and regulated by a common operator. It is defined as a set of adjacent structural genes, plus the adjacent regulatory signals that affect transcription of the structural genes.<sup>5</sup><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> The regulators of a given operon, including <a href="/wiki/Repressor" title="Repressor">repressors</a>, <a href="/wiki/Corepressor_(genetics)" class="mw-redirect" title="Corepressor (genetics)">corepressors</a>, and <a href="/wiki/Activator_(genetics)" title="Activator (genetics)">activators</a>, are not necessarily coded for by that operon. The location and condition of the regulators, promoter, operator and structural DNA sequences can determine the effects of common mutations. </p><p>Operons are related to <a href="/wiki/Regulon" title="Regulon">regulons</a>, <a href="/wiki/Stimulon" class="mw-redirect" title="Stimulon">stimulons</a> and <a href="/wiki/Modulon" class="mw-redirect" title="Modulon">modulons</a>; whereas operons contain a set of genes regulated by the same operator, regulons contain a set of genes under regulation by a single regulatory protein, and stimulons contain a set of genes under regulation by a single cell stimulus. According to its authors, the term "operon" is derived from the verb "to operate".<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="As_a_unit_of_transcription">As a unit of transcription</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=3" title="Edit section: As a unit of transcription"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An operon contains one or more <a href="/wiki/Structural_gene" title="Structural gene">structural genes</a> which are generally transcribed into one <a href="/wiki/Polycistronic" class="mw-redirect" title="Polycistronic">polycistronic</a> <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> (a single mRNA molecule that codes for more than one <a href="/wiki/Protein" title="Protein">protein</a>). However, the definition of an operon does not require the mRNA to be polycistronic, though in practice, it usually is.<sup id="cite_ref-Operons_in_eukaryotes_6-1" class="reference"><a href="#cite_note-Operons_in_eukaryotes-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Upstream of the structural genes lies a <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a> sequence which provides a site for <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> to bind and initiate transcription. Close to the promoter lies a section of DNA called an <i>operator</i>. </p> <div class="mw-heading mw-heading2"><h2 id="Operons_versus_clustering_of_prokaryotic_genes">Operons versus clustering of prokaryotic genes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=4" title="Edit section: Operons versus clustering of prokaryotic genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>All the <a href="/wiki/Structural_gene" title="Structural gene">structural genes</a> of an operon are turned ON or OFF together, due to a single promoter and operator upstream to them, but sometimes more control over the gene expression is needed. To achieve this aspect, some bacterial genes are located near together, but there is a specific promoter for each of them; this is called <a href="/wiki/Metabolic_gene_cluster" title="Metabolic gene cluster">gene clustering</a>. Usually these genes encode proteins which will work together in the same pathway, such as a metabolic pathway. Gene clustering helps a prokaryotic cell to produce metabolic enzymes in a correct order.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> In one study, it has been posited that in the <a href="/wiki/Asgard_(archaea)" class="mw-redirect" title="Asgard (archaea)">Asgard (archaea)</a>, ribosomal protein coding genes occur in clusters that are less conserved in their organization than in other <a href="/wiki/Archaea" title="Archaea">Archaea</a>; the closer an <a href="/wiki/Asgard_(archaea)" class="mw-redirect" title="Asgard (archaea)">Asgard (archaea)</a> is to the <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, the more dispersed is the arrangement of the ribosomal protein coding genes.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="General_structure">General structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=5" title="Edit section: General structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><span class="anchor" id="General_structure"></span> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Lac_Operon.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Lac_Operon.svg/300px-Lac_Operon.svg.png" decoding="async" width="300" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Lac_Operon.svg/450px-Lac_Operon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/22/Lac_Operon.svg/600px-Lac_Operon.svg.png 2x" data-file-width="512" data-file-height="382" /></a><figcaption><b><i>1</i>: RNA Polymerase, <i>2</i>: Repressor, <i>3</i>: Promoter, <i>4</i>: Operator, <i>5</i>: Lactose, <i>6</i>: lacZ, <i>7</i>: lacY, <i>8</i>: lacA. </b> <b>Top</b>: The gene is essentially turned off. There is no lactose to inhibit the repressor, so the repressor binds to the operator, which obstructs the RNA polymerase from binding to the promoter and making lactase. <b>Bottom</b>: The gene is turned on. Lactose is inhibiting the repressor, allowing the RNA polymerase to bind with the promoter, and express the genes, which synthesize lactase. Eventually, the lactase will digest all of the lactose, until there is none to bind to the repressor. The repressor will then bind to the operator, stopping the manufacture of lactase.</figcaption></figure> <p>An operon is made up of 3 basic DNA components: </p> <ul><li><a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">Promoter</a> – a <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> sequence that enables a gene to be <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a>. The promoter is recognized by <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>, which then initiates transcription. In RNA synthesis, promoters indicate which genes should be used for messenger RNA creation – and, by extension, control which proteins the cell produces.</li> <li><span class="anchor" id="Operator"></span> <b>Operator</b> – a segment of <a href="/wiki/DNA" title="DNA">DNA</a> to which a <a href="/wiki/Repressor" title="Repressor">repressor</a> binds. It is classically defined in the <a href="/wiki/Lac_operon" title="Lac operon">lac operon</a> as a segment between the promoter and the genes of the operon.<sup id="cite_ref-blewin_16-0" class="reference"><a href="#cite_note-blewin-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The main operator (O1) in the <i>lac</i> operon is located slightly downstream of the promoter; two additional operators, O2 and O3 are located at -82 and +412, respectively. In the case of a repressor, the repressor protein physically obstructs the RNA polymerase from transcribing the genes.</li> <li><a href="/wiki/Structural_gene" title="Structural gene">Structural genes</a> – the genes that are co-regulated by the operon.</li></ul> <p>Not always included within the operon, but important in its function is a <a href="/wiki/Regulatory_gene" class="mw-redirect" title="Regulatory gene">regulatory gene</a>, a constantly expressed gene which codes for <a href="/wiki/Repressor_proteins" class="mw-redirect" title="Repressor proteins">repressor proteins</a>. The regulatory gene does not need to be in, adjacent to, or even near the operon to control it.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>An <a href="/wiki/Inducer" title="Inducer">inducer</a> (small molecule) can displace a repressor (protein) from the operator site (DNA), resulting in an uninhibited operon. </p><p>Alternatively, a <a href="/wiki/Corepressor" title="Corepressor">corepressor</a> can bind to the repressor to allow its binding to the operator site. A good example of this type of regulation is seen for the <a href="/wiki/Trp_operon" title="Trp operon">trp operon</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Regulation">Regulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=6" title="Edit section: Regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Control of an operon is a type of <a href="/wiki/Gene_regulation" class="mw-redirect" title="Gene regulation">gene regulation</a> that enables organisms to regulate the expression of various genes depending on environmental conditions. Operon regulation can be either negative or positive by induction or repression.<sup id="cite_ref-blewin_16-1" class="reference"><a href="#cite_note-blewin-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Negative control involves the binding of a <a href="/wiki/Repressor" title="Repressor">repressor</a> to the operator to prevent transcription. </p> <ul><li>In <i>negative inducible operons</i>, a regulatory repressor protein is normally bound to the operator, which prevents the transcription of the genes on the operon. If an <a href="/wiki/Inducer" title="Inducer">inducer</a> molecule is present, it binds to the repressor and changes its conformation so that it is unable to bind to the operator. This allows for expression of the operon. The <a href="/wiki/Lac_operon" title="Lac operon"><i>lac</i> operon</a> is a negatively controlled inducible operon, where the inducer molecule is <a href="/wiki/Allolactose" title="Allolactose">allolactose</a>.</li> <li>In <i>negative repressible operons</i>, transcription of the operon normally takes place. Repressor proteins are produced by a <a href="/wiki/Regulator_gene" title="Regulator gene">regulator gene</a>, but they are unable to bind to the operator in their normal conformation. However, certain molecules called corepressors are bound by the repressor protein, causing a conformational change to the active site. The activated repressor protein binds to the operator and prevents transcription. The <a href="/wiki/Trp_operon" title="Trp operon"><i>trp</i> operon</a>, involved in the synthesis of <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> (which itself acts as the corepressor), is a negatively controlled repressible operon.</li></ul> <p>Operons can also be positively controlled. With positive control, an <a href="/wiki/Activator_(genetics)" title="Activator (genetics)">activator</a> protein stimulates transcription by binding to DNA (usually at a site other than the operator). </p> <ul><li>In <i>positive inducible operons</i>, activator proteins are normally unable to bind to the pertinent DNA. When an <a href="/wiki/Inducer" title="Inducer">inducer</a> is bound by the activator protein, it undergoes a change in conformation so that it can bind to the DNA and activate transcription. Examples of positive inducible operons include the MerR family of transcriptional activators.</li> <li>In <i>positive repressible operons</i>, the activator proteins are normally bound to the pertinent DNA segment. However, when an <a href="/wiki/Reaction_inhibitor" title="Reaction inhibitor">inhibitor</a> is bound by the activator, it is prevented from binding the DNA. This stops activation and transcription of the system.</li></ul> <div class="mw-heading mw-heading2"><h2 id="The_lac_operon">The <i>lac</i> operon</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=7" title="Edit section: The lac operon"><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/Lac_operon" title="Lac operon">lac operon</a></div> <p>The <i>lac</i> operon of the <a href="/wiki/Model_organism" title="Model organism">model</a> bacterium <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i> was the first operon to be discovered and provides a typical example of operon function. It consists of three adjacent <a href="/wiki/Structural_gene" title="Structural gene">structural genes</a>, a <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a>, a <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">terminator</a>, and an <a href="/wiki/Operator_(biology)" class="mw-redirect" title="Operator (biology)">operator</a>. The <i>lac</i> operon is regulated by several factors including the availability of <a href="/wiki/Glucose" title="Glucose">glucose</a> and <a href="/wiki/Lactose" title="Lactose">lactose</a>. It can be activated by <a href="/wiki/Allolactose" title="Allolactose">allolactose</a>. Lactose binds to the repressor protein and prevents it from repressing gene transcription. This is an example of the <a href="/wiki/Derepression" title="Derepression">derepressible</a> (from above: negative inducible) model. So it is a negative inducible operon induced by presence of lactose or allolactose. </p> <div class="mw-heading mw-heading2"><h2 id="The_trp_operon">The <i>trp</i> operon</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=8" title="Edit section: The trp operon"><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/Trp_operon" title="Trp operon">trp operon</a></div> <p>Discovered in 1953 by <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> and colleagues, the trp operon in <i>E. coli</i> was the first repressible operon to be discovered. While the lac operon can be activated by a chemical (<a href="/wiki/Allolactose" title="Allolactose">allolactose</a>), the tryptophan (Trp) operon is inhibited by a chemical (tryptophan). This operon contains five structural genes: trp E, trp D, trp C, trp B, and trp A, which encodes <a href="/wiki/Tryptophan_synthetase" class="mw-redirect" title="Tryptophan synthetase">tryptophan synthetase</a>. It also contains a promoter which binds to RNA polymerase and an operator which blocks transcription when bound to the protein synthesized by the repressor gene (trp R) that binds to the operator. In the lac operon, lactose binds to the repressor protein and prevents it from repressing gene transcription, while in the trp operon, tryptophan binds to the repressor protein and enables it to repress gene transcription. Also unlike the lac operon, the trp operon contains a leader peptide and an <a href="/wiki/Attenuator_(genetics)" title="Attenuator (genetics)">attenuator</a> sequence which allows for graded regulation.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> This is an example of the <a href="/wiki/Corepressor_(genetics)" class="mw-redirect" title="Corepressor (genetics)">corepressible</a> model. </p> <div class="mw-heading mw-heading2"><h2 id="Predicting_the_number_and_organization_of_operons">Predicting the number and organization of operons</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=9" title="Edit section: Predicting the number and organization of operons"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The number and organization of operons has been studied most critically in <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a></i>. As a result, predictions can be made based on an organism's genomic sequence. </p><p>One prediction method uses the intergenic distance between reading frames as a primary predictor of the number of operons in the genome. The separation merely changes the frame and guarantees that the read through is efficient. Longer stretches exist where operons start and stop, often up to 40–50 bases.<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> </p><p>An alternative method to predict operons is based on finding gene clusters where gene order and orientation is conserved in two or more genomes.<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> </p><p>Operon prediction is even more accurate if the functional class of the molecules is considered. Bacteria have clustered their reading frames into units, sequestered by co-involvement in protein complexes, common pathways, or shared substrates and transporters. Thus, accurate prediction would involve all of these data, a difficult task indeed. </p><p><a href="/wiki/Pascale_Cossart" title="Pascale Cossart">Pascale Cossart</a>'s laboratory was the first to experimentally identify all operons of a microorganism, <i><a href="/wiki/Listeria_monocytogenes" title="Listeria monocytogenes">Listeria monocytogenes</a></i>. The 517 polycistronic operons are listed in a 2009 study describing the global changes in transcription that occur in <i>L. monocytogenes</i> under different conditions.<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> <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=Operon&action=edit&section=10" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239009302">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output 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class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Operon&action=edit&section=11" 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-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</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 q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFSadavaHillisHellerBerenbaum2009" class="citation book cs1">Sadava DE, Hillis DM, Heller HC, Berenbaum M (2009). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=ANT8VB14oBUC&pg=PA349"><i>Life: The Science of Biology</i></a> (9th ed.). Macmillan. p. 349. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4292-1962-4" title="Special:BookSources/978-1-4292-1962-4"><bdi>978-1-4292-1962-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Life%3A+The+Science+of+Biology&rft.pages=349&rft.edition=9th&rft.pub=Macmillan&rft.date=2009&rft.isbn=978-1-4292-1962-4&rft.aulast=Sadava&rft.aufirst=David+E.&rft.au=Hillis%2C+David+M.&rft.au=Heller%2C+H.+Craig&rft.au=Berenbaum%2C+May&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DANT8VB14oBUC%26pg%3DPA349&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLodishZipurskyMatsudairaBaltimore2000" class="citation book cs1">Lodish H, Zipursky L, Matsudaira P, Baltimore D, Darnel J (2000). 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Freeman. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7167-3136-8" title="Special:BookSources/978-0-7167-3136-8"><bdi>978-0-7167-3136-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+9%3A+Molecular+Definition+of+a+Gene&rft.btitle=Molecular+Cell+Biology&rft.pub=W.+H.+Freeman&rft.date=2000&rft.isbn=978-0-7167-3136-8&rft.aulast=Lodish&rft.aufirst=Harvey&rft.au=Zipursky%2C+Lawrence&rft.au=Matsudaira%2C+Paul&rft.au=Baltimore%2C+David&rft.au=Darnel%2C+James&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmolecularcellbio00lodi&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-Kominek-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kominek_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kominek_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKominekDoeringOpulenteShen2019" class="citation journal cs1">Kominek J, Doering DT, Opulente DA, Shen XX, Zhou X, DeVirgilio J, Hulfachor AB, Groenewald M, Mcgee MA, Karlen SD, Kurtzman CP, Rokas A, Hittinger CT (March 2019). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7295392">"Eukaryotic Acquisition of a Bacterial Operon"</a>. <i>Cell</i>. <b>176</b> (6): 1356–1366.e10. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cell.2019.01.034">10.1016/j.cell.2019.01.034</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/PMC7295392">7295392</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/30799038">30799038</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=Eukaryotic+Acquisition+of+a+Bacterial+Operon&rft.volume=176&rft.issue=6&rft.pages=1356-1366.e10&rft.date=2019-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7295392%23id-name%3DPMC&rft_id=info%3Apmid%2F30799038&rft_id=info%3Adoi%2F10.1016%2Fj.cell.2019.01.034&rft.aulast=Kominek&rft.aufirst=J&rft.au=Doering%2C+DT&rft.au=Opulente%2C+DA&rft.au=Shen%2C+XX&rft.au=Zhou%2C+X&rft.au=DeVirgilio%2C+J&rft.au=Hulfachor%2C+AB&rft.au=Groenewald%2C+M&rft.au=Mcgee%2C+MA&rft.au=Karlen%2C+SD&rft.au=Kurtzman%2C+CP&rft.au=Rokas%2C+A&rft.au=Hittinger%2C+CT&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7295392&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSpiethBrookeKuerstenLea1993" class="citation journal cs1">Spieth J, Brooke G, Kuersten S, Lea K, Blumenthal T (May 1993). 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=Operons+in+C.+elegans%3A+polycistronic+mRNA+precursors+are+processed+by+trans-splicing+of+SL2+to+downstream+coding+regions&rft.volume=73&rft.issue=3&rft.pages=521-32&rft.date=1993-05&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A26918553%23id-name%3DS2CID&rft_id=info%3Apmid%2F8098272&rft_id=info%3Adoi%2F10.1016%2F0092-8674%2893%2990139-H&rft.aulast=Spieth&rft.aufirst=J&rft.au=Brooke%2C+G&rft.au=Kuersten%2C+S&rft.au=Lea%2C+K&rft.au=Blumenthal%2C+T&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrognaAshburner1997" class="citation journal cs1">Brogna S, Ashburner M (April 1997). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1169805">"The Adh-related gene of Drosophila melanogaster is expressed as a functional dicistronic messenger RNA: multigenic transcription in higher organisms"</a>. <i>The EMBO Journal</i>. <b>16</b> (8): 2023–31. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Femboj%2F16.8.2023">10.1093/emboj/16.8.2023</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/PMC1169805">1169805</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/9155028">9155028</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+EMBO+Journal&rft.atitle=The+Adh-related+gene+of+Drosophila+melanogaster+is+expressed+as+a+functional+dicistronic+messenger+RNA%3A+multigenic+transcription+in+higher+organisms&rft.volume=16&rft.issue=8&rft.pages=2023-31&rft.date=1997-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1169805%23id-name%3DPMC&rft_id=info%3Apmid%2F9155028&rft_id=info%3Adoi%2F10.1093%2Femboj%2F16.8.2023&rft.aulast=Brogna&rft.aufirst=S&rft.au=Ashburner%2C+M&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1169805&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-Operons_in_eukaryotes-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Operons_in_eukaryotes_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Operons_in_eukaryotes_6-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBlumenthal2004" class="citation journal cs1">Blumenthal T (November 2004). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fbfgp%2F3.3.199">"Operons in eukaryotes"</a>. <i>Briefings in Functional Genomics & Proteomics</i>. <b>3</b> (3): 199–211. <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/10.1093%2Fbfgp%2F3.3.199">10.1093/bfgp/3.3.199</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/15642184">15642184</a>.</cite><span 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text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC18690">18690</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/10823905">10823905</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=Operons+in+Escherichia+coli%3A+genomic+analyses+and+predictions&rft.volume=97&rft.issue=12&rft.pages=6652-7&rft.date=2000-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC18690%23id-name%3DPMC&rft_id=info%3Apmid%2F10823905&rft_id=info%3Adoi%2F10.1073%2Fpnas.110147297&rft_id=info%3Abibcode%2F2000PNAS...97.6652S&rft.aulast=Salgado&rft.aufirst=H&rft.au=Moreno-Hagelsieb%2C+G&rft.au=Smith%2C+TF&rft.au=Collado-Vides%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC18690&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFErmolaevaWhiteSalzberg2001" class="citation journal cs1">Ermolaeva MD, White O, Salzberg SL (March 2001). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC29727">"Prediction of operons in microbial genomes"</a>. <i>Nucleic Acids Research</i>. <b>29</b> (5): 1216–21. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fnar%2F29.5.1216">10.1093/nar/29.5.1216</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/PMC29727">29727</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/11222772">11222772</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nucleic+Acids+Research&rft.atitle=Prediction+of+operons+in+microbial+genomes&rft.volume=29&rft.issue=5&rft.pages=1216-21&rft.date=2001-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC29727%23id-name%3DPMC&rft_id=info%3Apmid%2F11222772&rft_id=info%3Adoi%2F10.1093%2Fnar%2F29.5.1216&rft.aulast=Ermolaeva&rft.aufirst=MD&rft.au=White%2C+O&rft.au=Salzberg%2C+SL&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC29727&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFToledo-AranaDussurgetNikitasSesto2009" class="citation journal cs1">Toledo-Arana A, Dussurget O, Nikitas G, Sesto N, Guet-Revillet H, Balestrino D, Loh E, Gripenland J, Tiensuu T, Vaitkevicius K, Barthelemy M, Vergassola M, Nahori MA, Soubigou G, Régnault B, Coppée JY, Lecuit M, Johansson J, Cossart P (June 2009). "The Listeria transcriptional landscape from saprophytism to virulence". <i>Nature</i>. <b>459</b> (7249): 950–6. <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/2009Natur.459..950T">2009Natur.459..950T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature08080">10.1038/nature08080</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19448609">19448609</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4341657">4341657</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=The+Listeria+transcriptional+landscape+from+saprophytism+to+virulence&rft.volume=459&rft.issue=7249&rft.pages=950-6&rft.date=2009-06&rft_id=info%3Adoi%2F10.1038%2Fnature08080&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4341657%23id-name%3DS2CID&rft_id=info%3Apmid%2F19448609&rft_id=info%3Abibcode%2F2009Natur.459..950T&rft.aulast=Toledo-Arana&rft.aufirst=A&rft.au=Dussurget%2C+O&rft.au=Nikitas%2C+G&rft.au=Sesto%2C+N&rft.au=Guet-Revillet%2C+H&rft.au=Balestrino%2C+D&rft.au=Loh%2C+E&rft.au=Gripenland%2C+J&rft.au=Tiensuu%2C+T&rft.au=Vaitkevicius%2C+K&rft.au=Barthelemy%2C+M&rft.au=Vergassola%2C+M&rft.au=Nahori%2C+MA&rft.au=Soubigou%2C+G&rft.au=R%C3%A9gnault%2C+B&rft.au=Copp%C3%A9e%2C+JY&rft.au=Lecuit%2C+M&rft.au=Johansson%2C+J&rft.au=Cossart%2C+P&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOperon" class="Z3988"></span></span> </li> </ol></div></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=Operon&action=edit&section=12" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://archive.today/20121214195153/http://www.broad.mit.edu/annotation/tbdb/operon/"><i>Mycobacterium tuberculosis</i> H37Rv Operon Correlation Browser</a></li> <li><a rel="nofollow" class="external text" href="http://operondb.jp">OBD</a> - <a href="/wiki/Operon_database" title="Operon database">Operon database</a> (a bit awkward to use though)</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 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td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Transcription_(Bacterial,_Eukaryotic)" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><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 abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Transcription" title="Template:Transcription"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Transcription" title="Template talk:Transcription"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Transcription" title="Special:EditPage/Template:Transcription"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Transcription_(Bacterial,_Eukaryotic)" style="font-size:114%;margin:0 4em"><a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription</a> (<a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a>, <a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional regulation</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">prokaryotic</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">Operon</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Trp_operon" title="Trp operon">trp operon</a></li> <li><a href="/wiki/Gab_operon" title="Gab operon">gab operon</a></li> <li><a href="/wiki/Gua_Operon" title="Gua Operon">Gua Operon</a></li> <li><a href="/wiki/L-arabinose_operon" title="L-arabinose operon">ara operon</a></li> <li><a href="/wiki/Gal_operon" title="Gal operon">gal operon</a></li></ul></li> <li><a href="/wiki/Repressor" title="Repressor">Repressor</a> <ul><li><a href="/wiki/Lac_repressor" title="Lac repressor">lac repressor</a></li> <li><a href="/wiki/Tryptophan_repressor" title="Tryptophan repressor">trp repressor</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">eukaryotic</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/Histone-modifying_enzymes" title="Histone-modifying enzymes">Histone-modifying enzymes</a><br />(<a href="/wiki/Histone" title="Histone">histone</a>/<a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</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/Histone_methylation" title="Histone methylation">Histone methylation</a>/<a href="/wiki/Histone_methyltransferase" title="Histone methyltransferase">Histone methyltransferase</a> <ul><li><a href="/wiki/EZH2" title="EZH2">EZH2</a></li></ul></li> <li><a href="/wiki/Demethylase" title="Demethylase">Histone demethylase</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</a> <ul><li><a href="/wiki/Histone_deacetylase" title="Histone deacetylase">Histone deacetylase</a> <a href="/wiki/HDAC1" title="HDAC1">HDAC1</a></li> <li><a href="/wiki/Histone_acetyltransferase" title="Histone acetyltransferase">Histone acetyltransferase</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</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><ul><li><a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferase</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Chromatin_remodeling" title="Chromatin remodeling">Chromatin remodeling</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/CHD7" title="CHD7">CHD7</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">both</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/Transcription_coregulator" title="Transcription coregulator">Transcription coregulator</a> <ul><li><a href="/wiki/Activator_(genetics)" title="Activator (genetics)">Activator</a></li> <li><a href="/wiki/Coactivator_(genetics)" title="Coactivator (genetics)">Coactivator</a></li> <li><a href="/wiki/Corepressor" title="Corepressor">Corepressor</a></li></ul></li> <li><a href="/wiki/Inducer" title="Inducer">Inducer</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Promotion</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> <ul><li><a href="/wiki/Pribnow_box" title="Pribnow box">Pribnow box</a></li> <li><a href="/wiki/TATA_box" title="TATA box">TATA box</a></li> <li><a href="/wiki/B_recognition_element" title="B recognition element">BRE</a></li> <li><a href="/wiki/CAAT_box" title="CAAT box">CAAT box</a></li> <li><a href="/wiki/Response_element" title="Response element">Response element</a></li></ul></li> <li><a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> <ul><li><a href="/wiki/E-box" title="E-box">E-box</a></li> <li><a href="/wiki/Response_element" title="Response element">Response element</a></li></ul></li> <li><a href="/wiki/Insulator_(genetics)" title="Insulator (genetics)">Insulator</a></li> <li><a href="/wiki/Silencer_(DNA)" class="mw-redirect" title="Silencer (DNA)">Silencer</a></li> <li><a href="/wiki/Internal_control_region" title="Internal control region">Internal control region</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Initiation</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription#Initiation" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Eukaryotic_transcription#Initiation" title="Eukaryotic transcription">Eukaryotic</a></li> <li><a href="/wiki/Archaeal_transcription_factor_B" title="Archaeal transcription factor B">Archaeal transcription factor B</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elongation</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription#Elongation" title="Bacterial transcription">bacterial</a> <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>: <a href="/wiki/RpoB" title="RpoB">rpoB</a></li> <li><a href="/wiki/Eukaryotic_transcription#Transcription_process" title="Eukaryotic transcription">eukaryotic</a> <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>: <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Termination<br />(<a href="/wiki/Bacterial_transcription#Termination" title="Bacterial transcription">bacterial</a>,<br /> <a href="/wiki/Eukaryotic_transcription#Termination" title="Eukaryotic transcription">eukaryotic</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">Terminator</a></li> <li><a href="/wiki/Intrinsic_termination" title="Intrinsic termination">Intrinsic termination</a></li> <li><a href="/wiki/Rho_factor" title="Rho factor">Rho factor</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_expression" 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_expression" 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 href="/wiki/Genetic_code" title="Genetic code">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"></div><div role="navigation" class="navbox" aria-labelledby="The_development_of_phenotype" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><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:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a class="mw-selflink selflink">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Systems</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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</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/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a href="/wiki/Morphogen" title="Morphogen">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><i><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></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"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a>: National <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q139677#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4210146-3">Germany</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Operons"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85095032">United States</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007548424205171">Israel</a></span></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Operon&oldid=1220963498">https://en.wikipedia.org/w/index.php?title=Operon&oldid=1220963498</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Gene_expression" title="Category:Gene expression">Gene expression</a></li><li><a href="/wiki/Category:Operons" title="Category:Operons">Operons</a></li></ul></div><div id="mw-hidden-catlinks" class="mw-hidden-catlinks mw-hidden-cats-hidden">Hidden categories: <ul><li><a href="/wiki/Category:CS1_French-language_sources_(fr)" title="Category:CS1 French-language sources (fr)">CS1 French-language sources (fr)</a></li><li><a href="/wiki/Category:Articles_with_short_description" title="Category:Articles with short description">Articles with short description</a></li><li><a href="/wiki/Category:Short_description_matches_Wikidata" title="Category:Short description matches Wikidata">Short description matches Wikidata</a></li></ul></div></div> </div> </main> </div> <div class="mw-footer-container"> <footer id="footer" class="mw-footer" > <ul id="footer-info"> <li id="footer-info-lastmod"> This page was last edited on 27 April 2024, at 00:40<span class="anonymous-show"> (UTC)</span>.</li> <li id="footer-info-copyright">Text is available under the <a href="/wiki/Wikipedia:Text_of_the_Creative_Commons_Attribution-ShareAlike_4.0_International_License" title="Wikipedia:Text of the Creative Commons Attribution-ShareAlike 4.0 International License">Creative Commons Attribution-ShareAlike 4.0 License</a>; additional terms may apply. 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