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class="vector-toc-numb">2</span> <span>Structure and function</span> </div> </a> <button aria-controls="toc-Structure_and_function-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Structure and function subsection</span> </button> <ul id="toc-Structure_and_function-sublist" class="vector-toc-list"> <li id="toc-F1_region" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#F1_region"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>F<sub>1</sub> region</span> </div> </a> <ul id="toc-F1_region-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-FO_region" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#FO_region"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>F<sub>O</sub> region</span> </div> </a> <ul id="toc-FO_region-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Binding_model" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Binding_model"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Binding model</span> </div> </a> <ul id="toc-Binding_model-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Physiological_role" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Physiological_role"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Physiological role</span> </div> </a> <ul id="toc-Physiological_role-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inhibitors" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Inhibitors"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Inhibitors</span> </div> </a> <ul id="toc-Inhibitors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_different_organisms" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#In_different_organisms"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>In different organisms</span> </div> </a> <button aria-controls="toc-In_different_organisms-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle In different organisms subsection</span> </button> <ul id="toc-In_different_organisms-sublist" class="vector-toc-list"> <li id="toc-Bacteria" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bacteria"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Bacteria</span> </div> </a> <ul id="toc-Bacteria-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Yeast" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Yeast"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Yeast</span> </div> </a> <ul id="toc-Yeast-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plant" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plant"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Plant</span> </div> </a> <ul id="toc-Plant-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mammal" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mammal"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>Mammal</span> </div> </a> <ul id="toc-Mammal-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_eukaryotes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Other_eukaryotes"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.5</span> <span>Other eukaryotes</span> </div> </a> <ul id="toc-Other_eukaryotes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Archaea" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Archaea"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.6</span> <span>Archaea</span> </div> </a> <ul id="toc-Archaea-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-LUCA_and_earlier" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#LUCA_and_earlier"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.7</span> <span>LUCA and earlier</span> </div> </a> <ul id="toc-LUCA_and_earlier-sublist" class="vector-toc-list"> </ul> </li> </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">8</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">9</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</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">ATP synthase</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 45 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-45" 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">45 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A3%D9%8A%D9%87_%D8%AA%D9%8A_%D8%A8%D9%8A_%D8%B3%D9%8A%D9%86%D8%AB%D8%A7%D8%B2" title="أيه تي بي سينثاز – Arabic" lang="ar" hreflang="ar" data-title="أيه تي بي سينثاز" 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%90%D0%A2%D0%A4_%D1%81%D0%B8%D0%BD%D1%82%D0%B0%D0%B7%D0%B0" title="АТФ синтаза – Bulgarian" lang="bg" hreflang="bg" data-title="АТФ синтаза" 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/ATP_sintaza" title="ATP sintaza – Bosnian" lang="bs" hreflang="bs" data-title="ATP sintaza" 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/ATP_sintetasa" title="ATP sintetasa – Catalan" lang="ca" hreflang="ca" data-title="ATP sintetasa" 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/ATP_synt%C3%A1za" title="ATP syntáza – Czech" lang="cs" hreflang="cs" data-title="ATP syntáza" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/ATP-syntase" title="ATP-syntase – Danish" lang="da" hreflang="da" data-title="ATP-syntase" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://de.wikipedia.org/wiki/ATP-Synthase" title="ATP-Synthase – German" lang="de" hreflang="de" data-title="ATP-Synthase" 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/ATP_s%C3%BCntaas" title="ATP süntaas – Estonian" lang="et" hreflang="et" data-title="ATP süntaas" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/ATP_sintasa" title="ATP sintasa – Spanish" lang="es" hreflang="es" data-title="ATP sintasa" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/ATP_sintasa" title="ATP sintasa – Basque" lang="eu" hreflang="eu" data-title="ATP sintasa" 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%DB%8C%E2%80%8C%D8%AA%DB%8C%E2%80%8C%D9%BE%DB%8C_%D8%B3%D9%86%D8%AA%D8%A7%D8%B2" title="ای‌تی‌پی سنتاز – Persian" lang="fa" hreflang="fa" data-title="ای‌تی‌پی سنتاز" 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/ATP_synthase" title="ATP synthase – French" lang="fr" hreflang="fr" data-title="ATP synthase" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/ATP_sintase" title="ATP sintase – Galician" lang="gl" hreflang="gl" data-title="ATP sintase" 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/ATP_%EC%83%9D%EC%84%B1%ED%9A%A8%EC%86%8C" title="ATP 생성효소 – Korean" lang="ko" hreflang="ko" data-title="ATP 생성효소" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B1%D4%B5%D5%96-%D5%BD%D5%AB%D5%B6%D5%A9%D5%A1%D5%A6" title="ԱԵՖ-սինթազ – Armenian" lang="hy" hreflang="hy" data-title="ԱԵՖ-սինթազ" 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/ATP_sintaza" title="ATP sintaza – Croatian" lang="hr" hreflang="hr" data-title="ATP sintaza" 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/ATP_sintase" title="ATP sintase – Indonesian" lang="id" hreflang="id" data-title="ATP sintase" 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/ATPasi_trasportante_H%2B_tra_due_settori" title="ATPasi trasportante H+ tra due settori – Italian" lang="it" hreflang="it" data-title="ATPasi trasportante H+ tra due settori" 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/ATP_%D7%A1%D7%99%D7%A0%D7%AA%D7%90%D7%96" title="ATP סינתאז – Hebrew" lang="he" hreflang="he" data-title="ATP סינתאז" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Sintase_ATP" title="Sintase ATP – Javanese" lang="jv" hreflang="jv" data-title="Sintase ATP" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/ATP_sentaz" title="ATP sentaz – Kurdish" lang="ku" hreflang="ku" data-title="ATP sentaz" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/ATP_synthasis" title="ATP synthasis – Latin" lang="la" hreflang="la" data-title="ATP synthasis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/ATP-szint%C3%A1z" title="ATP-szintáz – Hungarian" lang="hu" hreflang="hu" data-title="ATP-szintáz" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%90%D0%A2%D0%A4-%D1%81%D0%B8%D0%BD%D1%82%D0%B0%D0%B7%D0%B0" title="АТФ-синтаза – Macedonian" lang="mk" hreflang="mk" data-title="АТФ-синтаза" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/ATP_sintase" title="ATP sintase – Malay" lang="ms" hreflang="ms" data-title="ATP sintase" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/ATP-synthase" title="ATP-synthase – Dutch" lang="nl" hreflang="nl" data-title="ATP-synthase" 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/ATP%E5%90%88%E6%88%90%E9%85%B5%E7%B4%A0" title="ATP合成酵素 – Japanese" lang="ja" hreflang="ja" data-title="ATP合成酵素" 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-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/ATF-sintaza" title="ATF-sintaza – Uzbek" lang="uz" hreflang="uz" data-title="ATF-sintaza" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Syntaza_ATP" title="Syntaza ATP – Polish" lang="pl" hreflang="pl" data-title="Syntaza ATP" 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/ATP_sintase" title="ATP sintase – Portuguese" lang="pt" hreflang="pt" data-title="ATP sintase" 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/ATP_sintaz%C4%83" title="ATP sintază – Romanian" lang="ro" hreflang="ro" data-title="ATP sintază" 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%90%D0%A2%D0%A4-%D1%81%D0%B8%D0%BD%D1%82%D0%B0%D0%B7%D0%B0" title="АТФ-синтаза – Russian" lang="ru" hreflang="ru" data-title="АТФ-синтаза" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sc mw-list-item"><a href="https://sc.wikipedia.org/wiki/ATPasi_trasportanti_H%2B_in_mesu_a_d%C3%B9as_settoris" title="ATPasi trasportanti H+ in mesu a dùas settoris – Sardinian" lang="sc" hreflang="sc" data-title="ATPasi trasportanti H+ in mesu a dùas settoris" data-language-autonym="Sardu" data-language-local-name="Sardinian" class="interlanguage-link-target"><span>Sardu</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/ATP_synthase" title="ATP synthase – Simple English" lang="en-simple" hreflang="en-simple" data-title="ATP synthase" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/ATP_synt%C3%A1za" title="ATP syntáza – Slovak" lang="sk" hreflang="sk" data-title="ATP syntáza" 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/ATP-sintaza" title="ATP-sintaza – Slovenian" lang="sl" hreflang="sl" data-title="ATP-sintaza" 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/H%2B_transportuju%C4%87a_dvosektorna_ATPaza" title="H+ transportujuća dvosektorna ATPaza – Serbian" lang="sr" hreflang="sr" data-title="H+ transportujuća dvosektorna ATPaza" 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/H%2B_transportuju%C4%87a_dvosektorna_ATPaza" title="H+ transportujuća dvosektorna ATPaza – Serbo-Croatian" lang="sh" hreflang="sh" data-title="H+ transportujuća dvosektorna ATPaza" 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-su mw-list-item"><a href="https://su.wikipedia.org/wiki/ATP_sintase" title="ATP sintase – Sundanese" lang="su" hreflang="su" data-title="ATP sintase" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/ATP-syntaasi" title="ATP-syntaasi – Finnish" lang="fi" hreflang="fi" data-title="ATP-syntaasi" 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/ATP-syntas" title="ATP-syntas – Swedish" lang="sv" hreflang="sv" data-title="ATP-syntas" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/ATP_sentaz" title="ATP sentaz – Turkish" lang="tr" hreflang="tr" data-title="ATP sentaz" 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%90%D0%A2%D0%A4-%D1%81%D0%B8%D0%BD%D1%82%D0%B0%D0%B7%D0%B0" title="АТФ-синтаза – Ukrainian" lang="uk" hreflang="uk" data-title="АТФ-синтаза" 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/ATP_synthase" title="ATP synthase – Vietnamese" lang="vi" hreflang="vi" data-title="ATP synthase" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li 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The present address (URL) is a <a href="/wiki/Help:Permanent_link" title="Help:Permanent link">permanent link</a> to this version.</b></p><div id="revision-info-current-plain" style="display: none;">Revision as of 14:56, 17 September 2024 by <a href="/wiki/User:JeffSpaceman" class="mw-userlink" title="User:JeffSpaceman" data-mw-revid="1246202526"><bdi>JeffSpaceman</bdi></a> <span class="mw-usertoollinks">(<a href="/wiki/User_talk:JeffSpaceman" class="mw-usertoollinks-talk" title="User talk:JeffSpaceman">talk</a> | <a href="/wiki/Special:Contributions/JeffSpaceman" class="mw-usertoollinks-contribs" title="Special:Contributions/JeffSpaceman">contribs</a>)</span> <span class="comment">(Reverted edit by <a href="/wiki/Special:Contributions/104.152.208.62" title="Special:Contributions/104.152.208.62">104.152.208.62</a> (<a href="/wiki/User_talk:104.152.208.62" title="User talk:104.152.208.62">talk</a>) to last version by JeffSpaceman)</span></div></div><div id="mw-revision-nav">(<a href="/w/index.php?title=ATP_synthase&diff=prev&oldid=1246202526" title="ATP synthase">diff</a>) <a href="/w/index.php?title=ATP_synthase&direction=prev&oldid=1246202526" title="ATP synthase">← Previous revision</a> | Latest revision (diff) | Newer revision → (diff)</div></div></div></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Enzyme</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">ATP Synthase</th></tr><tr><td colspan="2" class="infobox-image"><span typeof="mw:File"><a href="/wiki/File:Atp_synthase.PNG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Atp_synthase.PNG/250px-Atp_synthase.PNG" decoding="async" width="250" height="333" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Atp_synthase.PNG/375px-Atp_synthase.PNG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/00/Atp_synthase.PNG/500px-Atp_synthase.PNG 2x" data-file-width="1200" data-file-height="1600" /></a></span><div class="infobox-caption">Molecular model of ATP synthase determined by <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>. Stator is not shown here.</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Enzyme_Commission_number" title="Enzyme Commission number">EC no.</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.enzyme-database.org/query.php?ec=7.1.2.2">7.1.2.2</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/CAS_registry_number" class="mw-redirect" title="CAS registry number">CAS no.</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://commonchemistry.cas.org/detail?cas_rn=9000-83-3&title=">9000-83-3 </a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Databases</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/IntEnz" title="IntEnz">IntEnz</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/intenz/query?cmd=SearchEC&ec=7.1.2.2">IntEnz view</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/BRENDA" title="BRENDA">BRENDA</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://www.brenda-enzymes.org/enzyme.php?ecno=7.1.2.2">BRENDA entry</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/ExPASy" class="mw-redirect" title="ExPASy">ExPASy</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://enzyme.expasy.org/EC/7.1.2.2">NiceZyme view</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/KEGG" title="KEGG">KEGG</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.genome.jp/dbget-bin/www_bget?enzyme+7.1.2.2">KEGG entry</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/MetaCyc" title="MetaCyc">MetaCyc</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://biocyc.org/META/substring-search?type=NIL&object=7.1.2.2">metabolic pathway</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/PRIAM_enzyme-specific_profiles" title="PRIAM enzyme-specific profiles">PRIAM</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://priam.prabi.fr/cgi-bin/PRIAM_profiles_CurrentRelease.pl?EC=7.1.2.2">profile</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> structures</th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity.rcsb_ec_lineage.id:7.1.2.2">RCSB PDB</a> <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?ec_number:7.1.2.2">PDBe</a> <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/enzymes/GetPage.pl?ec_number=7.1.2.2">PDBsum</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Gene_Ontology" title="Gene Ontology">Gene Ontology</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://amigo.geneontology.org/amigo/term/GO:0046961">AmiGO </a> / <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/QuickGO/term/GO:0046961">QuickGO</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Search</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PubMed_Central" title="PubMed Central">PMC</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&term=7.1.2.2%5BEC/RN%20Number%5D%20AND%20pubmed%20pmc%20local%5Bsb%5D">articles</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PubMed" title="PubMed">PubMed</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&term=7.1.2.2%5BEC/RN%20Number%5D">articles</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/protein?term=7.1.2.2%5BEC/RN%20Number%5D">proteins</a></td></tr></tbody></table></td></tr></tbody></table> <p><b>ATP synthase</b> is an <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> that catalyzes the formation of the energy storage molecule <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP) using <a href="/wiki/Adenosine_diphosphate" title="Adenosine diphosphate">adenosine diphosphate</a> (ADP) and inorganic <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> (P<sub>i</sub>). ATP synthase is a <a href="/wiki/Molecular_machine" title="Molecular machine">molecular machine</a>. The overall reaction catalyzed by ATP synthase is: </p> <ul><li>ADP + P<sub>i</sub> + 2H<sup>+</sup><sub>out</sub> ⇌ ATP + H<sub>2</sub>O + 2H<sup>+</sup><sub>in</sub></li></ul> <p>ATP synthase lies across a cellular membrane and forms an aperture that <a href="/wiki/Hydron_(chemistry)" title="Hydron (chemistry)">protons</a> can cross from areas of high concentration to areas of low concentration, imparting energy for the synthesis of ATP. This <a href="/wiki/Electrochemical_gradient" title="Electrochemical gradient">electrochemical gradient</a> is generated by the <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport chain</a> and allows cells to store energy in ATP for later use. In <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotic cells</a> ATP synthase lies across the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a>, while in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic cells</a> it lies across the <a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">inner mitochondrial membrane</a>. Organisms capable of <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a> also have ATP synthase across the <a href="/wiki/Thylakoid_membrane" class="mw-redirect" title="Thylakoid membrane">thylakoid membrane</a>, which in <a href="/wiki/Plants" class="mw-redirect" title="Plants">plants</a> is located in the <a href="/wiki/Chloroplast" title="Chloroplast">chloroplast</a> and in <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacteria</a> is located in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. </p><p>Eukaryotic ATP synthases are <a href="/wiki/F-ATPase" title="F-ATPase">F-ATPases</a>, running "in reverse" for an <a href="/wiki/ATPase" title="ATPase">ATPase</a>. This article deals mainly with this type. An F-ATPase consists of two main subunits, F<sub>O</sub> and F<sub>1</sub>, which has a rotational motor mechanism allowing for ATP production.<sup id="cite_ref-Okuno_2011_1-0" class="reference"><a href="#cite_note-Okuno_2011-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Junge_2015_2-0" class="reference"><a href="#cite_note-Junge_2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Nomenclature">Nomenclature</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=1" title="Edit section: Nomenclature"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The F<sub>1</sub> fraction derives its name from the term "Fraction 1" and F<sub>O</sub> (written as a subscript letter "o", not "zero") derives its name from being the binding fraction for <a href="/wiki/Oligomycin" title="Oligomycin">oligomycin</a>, a type of naturally derived antibiotic that is able to inhibit the F<sub>O</sub> unit of ATP synthase.<sup id="cite_ref-Kagawa_1966_3-0" class="reference"><a href="#cite_note-Kagawa_1966-3"><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> These functional regions consist of different protein subunits — refer to tables. This enzyme is used in synthesis of ATP through aerobic respiration. </p> <div class="mw-heading mw-heading2"><h2 id="Structure_and_function">Structure and function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=2" title="Edit section: Structure and function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/48/Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png/289px-Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png" decoding="async" width="289" height="178" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/48/Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png/434px-Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/48/Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png/578px-Fo_subunit_of_ATPase_C1._Picture_Created_in_PyMol.png 2x" data-file-width="657" data-file-height="404" /></a><figcaption>Bovine mitochondrial ATP synthase. The F<sub>O</sub>, F<sub>1</sub>, axle, and stator regions are color coded magenta, green, orange, and cyan respectively i.e. <span style="color:#f0f">F<sub>O</sub></span>, <span style="color:#0f0">F<sub>1</sub></span>, <span style="color:#f80">axle</span>, <span style="color:#0ff">stator</span>.<sup id="cite_ref-pmid26439008_5-0" class="reference"><a href="#cite_note-pmid26439008-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Atpsynthase.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Atpsynthase.jpg/150px-Atpsynthase.jpg" decoding="async" width="150" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Atpsynthase.jpg/225px-Atpsynthase.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/34/Atpsynthase.jpg/300px-Atpsynthase.jpg 2x" data-file-width="344" data-file-height="360" /></a><figcaption>Simplified model of F<sub>O</sub>F<sub>1</sub>-ATPase alias ATP synthase of <i>E. coli</i>. Subunits of the enzyme are labeled accordingly.</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Atpsyntase4.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Atpsyntase4.jpg/150px-Atpsyntase4.jpg" decoding="async" width="150" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Atpsyntase4.jpg/225px-Atpsyntase4.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/d/d1/Atpsyntase4.jpg 2x" data-file-width="265" data-file-height="333" /></a><figcaption>Rotation engine of ATP synthase.</figcaption></figure> <p>Located within the <a href="/wiki/Thylakoid_membrane" class="mw-redirect" title="Thylakoid membrane">thylakoid membrane</a> and the <a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">inner mitochondrial membrane</a>, ATP synthase consists of two regions F<sub>O</sub> and F<sub>1</sub>. F<sub>O</sub> causes rotation of F<sub>1</sub> and is made of c-ring and subunits a, two b, F6. F<sub>1</sub> is made of α, β, γ, and δ subunits. F<sub>1</sub> has a water-soluble part that can hydrolyze ATP. F<sub>O</sub> on the other hand has mainly hydrophobic regions. F<sub>O</sub> F<sub>1</sub> creates a pathway for protons movement across the membrane.<sup id="cite_ref-Velours_2000_7-0" class="reference"><a href="#cite_note-Velours_2000-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="F1_region">F<sub>1</sub> region</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=3" title="Edit section: F1 region"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The F<sub>1</sub> portion of ATP synthase is <a href="/wiki/Hydrophilic" class="mw-redirect" title="Hydrophilic">hydrophilic</a> and responsible for hydrolyzing ATP. The F<sub>1</sub> unit protrudes into the <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">mitochondrial matrix</a> space. Subunits α and β make a hexamer with 6 binding sites. Three of them are catalytically inactive and they bind ADP. </p><p>Three other subunits catalyze the ATP synthesis. The other F<sub>1</sub> subunits γ, δ, and ε are a part of a rotational motor mechanism (rotor/axle). The γ subunit allows β to go through conformational changes (i.e., closed, half open, and open states) that allow for ATP to be bound and released once synthesized. The F<sub>1</sub> particle is large and can be seen in the transmission electron microscope by negative staining.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> These are particles of 9 nm diameter that pepper the inner mitochondrial membrane. </p> <table class="wikitable" style="text-align:center"> <caption>F<sub>1</sub> – Subunits<sup id="cite_ref-pmid24878343_9-0" class="reference"><a href="#cite_note-pmid24878343-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th>Subunit</th> <th>Human Gene</th> <th>Note </th></tr> <tr> <td><a href="/wiki/ATP_synthase_alpha/beta_subunits" title="ATP synthase alpha/beta subunits">alpha</a></td> <td><i><a href="/wiki/ATP5A1" class="mw-redirect" title="ATP5A1">ATP5A1</a></i>, <i><a href="/wiki/ATPAF2" title="ATPAF2">ATPAF2</a></i></td> <td> </td></tr> <tr> <td><a href="/wiki/ATP_synthase_alpha/beta_subunits" title="ATP synthase alpha/beta subunits">beta</a></td> <td><i><a href="/wiki/ATP5B" title="ATP5B">ATP5B</a></i>, <i><a href="/wiki/ATPAF1" title="ATPAF1">ATPAF1</a></i></td> <td> </td></tr> <tr> <td><a href="/wiki/ATP_synthase_gamma_subunit" title="ATP synthase gamma subunit">gamma</a></td> <td><i><a href="/wiki/ATP5C1" title="ATP5C1">ATP5C1</a></i></td> <td> </td></tr> <tr> <td><a href="/wiki/ATP_synthase_delta/epsilon_subunit" title="ATP synthase delta/epsilon subunit">delta</a></td> <td><i><a href="/wiki/ATP5D" title="ATP5D">ATP5D</a></i></td> <td>Mitochondrial "delta" is bacterial/chloroplastic epsilon. </td></tr> <tr> <td><a href="/w/index.php?title=ATP_synthase_epsilon_subunit,_mitochondrial&action=edit&redlink=1" class="new" title="ATP synthase epsilon subunit, mitochondrial (page does not exist)">epsilon</a></td> <td><i><a href="/wiki/ATP5E" title="ATP5E">ATP5E</a></i></td> <td>Unique to mitochondria. </td></tr> <tr> <td><a href="/wiki/ATP_synthase_delta/OSCP_subunit" title="ATP synthase delta/OSCP subunit">OSCP</a></td> <td><i><a href="/wiki/ATP5O" class="mw-redirect" title="ATP5O">ATP5O</a></i></td> <td>Called "delta" in bacterial and chloroplastic versions. </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="FO_region">F<sub>O</sub> region</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=4" title="Edit section: FO region"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Fo_complex_subunit_F6.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Fo_complex_subunit_F6.png/285px-Fo_complex_subunit_F6.png" decoding="async" width="285" height="209" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Fo_complex_subunit_F6.png/428px-Fo_complex_subunit_F6.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Fo_complex_subunit_F6.png/570px-Fo_complex_subunit_F6.png 2x" data-file-width="661" data-file-height="485" /></a><figcaption>F<sub>O</sub> subunit F6 from the peripheral stalk region of ATP synthase.<sup id="cite_ref-Carbajo_2004_10-0" class="reference"><a href="#cite_note-Carbajo_2004-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>F<sub>O</sub> is a water <a href="/wiki/Insoluble" class="mw-redirect" title="Insoluble">insoluble</a> protein with eight subunits and a transmembrane ring. The ring has a <a href="/wiki/Tetramer" title="Tetramer">tetrameric</a> shape with a <a href="/wiki/Helix-loop-helix" class="mw-redirect" title="Helix-loop-helix">helix-loop-helix</a> protein that goes through conformational changes when protonated and deprotonated, pushing neighboring subunits to rotate, causing the spinning of F<sub>O</sub> which then also affects conformation of F<sub>1</sub>, resulting in switching of states of alpha and beta subunits. The F<sub>O</sub> region of ATP synthase is a proton pore that is embedded in the mitochondrial membrane. It consists of three main subunits, a, b, and c. Six c subunits make up the rotor ring, and subunit b makes up a stalk connecting to F<sub>1</sub> OSCP that prevents the αβ hexamer from rotating. Subunit a connects b to the c ring.<sup id="cite_ref-ecoli_11-0" class="reference"><a href="#cite_note-ecoli-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Humans have six additional subunits, <a href="/wiki/ATP5H" title="ATP5H">d</a>, <a href="/wiki/ATP5I" title="ATP5I">e</a>, <a href="/wiki/ATP5J2" title="ATP5J2">f</a>, <a href="/wiki/ATP5L" title="ATP5L">g</a>, <a href="/wiki/ATP5J" class="mw-redirect" title="ATP5J">F6</a>, and <a href="/wiki/MT-ATP8" title="MT-ATP8">8</a> (or A6L). This part of the enzyme is located in the mitochondrial inner membrane and couples proton translocation to the rotation that causes ATP synthesis in the F<sub>1</sub> region. </p><p>In eukaryotes, mitochondrial F<sub>O</sub> forms membrane-bending dimers. These dimers self-arrange into long rows at the end of the <a href="/wiki/Cristae" class="mw-redirect" title="Cristae">cristae</a>, possibly the first step of cristae formation.<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> An atomic model for the dimeric yeast F<sub>O</sub> region was determined by cryo-EM at an overall resolution of 3.6 Å.<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> <table class="wikitable" style="text-align:center"> <caption>F<sub>O</sub>-Main subunits </caption> <tbody><tr> <th>Subunit</th> <th>Human Gene </th></tr> <tr> <td>a</td> <td><i><a href="/wiki/MT-ATP6" title="MT-ATP6">MT-ATP6</a></i> </td></tr> <tr> <td>b</td> <td><i><a href="/wiki/ATP5PB" class="mw-redirect" title="ATP5PB">ATP5PB</a></i> </td></tr> <tr> <td><a href="/wiki/ATP_synthase_subunit_C" title="ATP synthase subunit C">c</a></td> <td><i><a href="/wiki/ATP5G1" title="ATP5G1">ATP5G1</a></i>, <i><a href="/wiki/ATP5G2" title="ATP5G2">ATP5G2</a></i>, <i><a href="/wiki/ATP5G3" title="ATP5G3">ATP5G3</a></i> </td></tr> </tbody></table> <div class="mw-heading mw-heading2"><h2 id="Binding_model">Binding model</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=5" title="Edit section: Binding model"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:ATPsyn.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/ATPsyn.gif/220px-ATPsyn.gif" decoding="async" width="220" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/6/62/ATPsyn.gif 1.5x" data-file-width="300" data-file-height="256" /></a><figcaption>Mechanism of ATP synthase. ADP and P<sub>i</sub> (pink) shown being combined into ATP (red), while the rotating γ (gamma) subunit in black causes conformational change.</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:ATP-Synthase.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/ATP-Synthase.svg/230px-ATP-Synthase.svg.png" decoding="async" width="230" height="402" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/ATP-Synthase.svg/345px-ATP-Synthase.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3d/ATP-Synthase.svg/460px-ATP-Synthase.svg.png 2x" data-file-width="512" data-file-height="894" /></a><figcaption>Depiction of ATP synthase using the <a href="/wiki/Chemiosmosis" title="Chemiosmosis">chemiosmotic</a> proton gradient to power ATP synthesis through <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a>.</figcaption></figure> <p>In the 1960s through the 1970s, <a href="/wiki/Paul_D._Boyer" title="Paul D. Boyer">Paul Boyer</a>, a <a href="/wiki/University_of_California,_Los_Angeles" title="University of California, Los Angeles">UCLA</a> Professor, developed the binding change, or flip-flop, mechanism theory, which postulated that ATP synthesis is dependent on a conformational change in ATP synthase generated by rotation of the gamma subunit. The research group of <a href="/wiki/John_E._Walker" title="John E. Walker">John E. Walker</a>, then at the <a href="/wiki/Laboratory_of_Molecular_Biology" class="mw-redirect" title="Laboratory of Molecular Biology">MRC Laboratory of Molecular Biology</a> in <a href="/wiki/Cambridge" title="Cambridge">Cambridge</a>, crystallized the F<sub>1</sub> catalytic-domain of ATP synthase. The structure, at the time the largest asymmetric protein structure known, indicated that Boyer's rotary-catalysis model was, in essence, correct. For elucidating this, Boyer and Walker shared half of the 1997 <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a>. </p><p>The crystal structure of the F<sub>1</sub> showed alternating alpha and beta <a href="/wiki/Protein_subunit" title="Protein subunit">subunits</a> (3 of each), arranged like segments of an orange around a rotating asymmetrical gamma subunit. According to the current model of ATP synthesis (known as the alternating catalytic model), the transmembrane potential created by (H+) proton cations supplied by the electron transport chain, drives the (H+) proton cations from the intermembrane space through the membrane via the F<sub>O</sub> region of ATP synthase. A portion of the F<sub>O</sub> (the ring of <a href="/wiki/ATP_synthase_subunit_C" title="ATP synthase subunit C">c-subunits</a>) <a href="/wiki/Rotating_locomotion_in_living_systems" title="Rotating locomotion in living systems">rotates</a> as the protons pass through the membrane. The <a href="/wiki/ATP_synthase_subunit_C" title="ATP synthase subunit C">c-ring</a> is tightly attached to the asymmetric central stalk (consisting primarily of the gamma subunit), causing it to rotate within the alpha<sub>3</sub>beta<sub>3</sub> of F<sub>1</sub> causing the 3 catalytic nucleotide binding sites to go through a series of conformational changes that lead to ATP synthesis. The major F<sub>1</sub> subunits are prevented from rotating in sympathy with the central stalk rotor by a peripheral stalk that joins the alpha<sub>3</sub>beta<sub>3</sub> to the non-rotating portion of F<sub>O</sub>. The structure of the intact ATP synthase is currently known at low-resolution from <a href="/wiki/Electron_cryo-microscopy" class="mw-redirect" title="Electron cryo-microscopy">electron cryo-microscopy</a> (cryo-EM) studies of the complex. The cryo-EM model of ATP synthase suggests that the peripheral stalk is a flexible structure that wraps around the complex as it joins F<sub>1</sub> to F<sub>O</sub>. Under the right conditions, the enzyme reaction can also be carried out in reverse, with ATP hydrolysis driving <a href="/wiki/Proton_pump" title="Proton pump">proton pumping</a> across the membrane. </p><p>The binding change mechanism involves the active site of a β subunit's cycling between three states.<sup id="cite_ref-Gresser_14-0" class="reference"><a href="#cite_note-Gresser-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> In the "loose" state, ADP and phosphate enter the active site; in the adjacent diagram, this is shown in pink. The enzyme then undergoes a change in shape and forces these molecules together, with the active site in the resulting "tight" state (shown in red) binding the newly produced ATP molecule with very high <a href="/wiki/Dissociation_constant" title="Dissociation constant">affinity</a>. Finally, the active site cycles back to the open state (orange), releasing ATP and binding more ADP and phosphate, ready for the next cycle of ATP production.<sup id="cite_ref-Nakamoto_15-0" class="reference"><a href="#cite_note-Nakamoto-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Physiological_role">Physiological role</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=6" title="Edit section: Physiological role"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Like other enzymes, the activity of F<sub>1</sub>F<sub>O</sub> ATP synthase is reversible. Large-enough quantities of ATP cause it to create a transmembrane <a href="/wiki/Proton" title="Proton">proton</a> <a href="/wiki/Gradient" title="Gradient">gradient</a>, this is used by fermenting bacteria that do not have an electron transport chain, but rather hydrolyze ATP to make a proton gradient, which they use to drive <a href="/wiki/Flagella" class="mw-redirect" title="Flagella">flagella</a> and the transport of nutrients into the cell. </p><p>In respiring <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> under physiological conditions, ATP synthase, in general, runs in the opposite direction, creating ATP while using the <a href="/wiki/Electrochemical_potential" title="Electrochemical potential">proton motive force</a> created by the <a href="/wiki/Electron_transfer_chain" class="mw-redirect" title="Electron transfer chain">electron transport chain</a> as a source of energy. The overall process of creating energy in this fashion is termed <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a>. The same process takes place in the <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a>, where ATP synthase is located in the inner mitochondrial membrane and the F<sub>1</sub>-part projects into the <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">mitochondrial matrix</a>. By pumping proton cations into the matrix, the ATP-synthase converts ADP into ATP. </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=7" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Evolution" title="Evolution">evolution</a> of ATP synthase is thought to have been modular whereby two functionally independent subunits became associated and gained new functionality.<sup id="cite_ref-RotaryDNA_16-0" class="reference"><a href="#cite_note-RotaryDNA-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-LifeScience_IL_17-0" class="reference"><a href="#cite_note-LifeScience_IL-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> This association appears to have occurred early in evolutionary history, because essentially the same structure and activity of ATP synthase enzymes are present in all kingdoms of life.<sup id="cite_ref-RotaryDNA_16-1" class="reference"><a href="#cite_note-RotaryDNA-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The F-ATP synthase displays high functional and mechanistic similarity to the <a href="/wiki/V-ATPase" title="V-ATPase">V-ATPase</a>.<sup id="cite_ref-InterPro_Synthase_18-0" class="reference"><a href="#cite_note-InterPro_Synthase-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> However, whereas the F-ATP synthase generates ATP by utilising a proton gradient, the <a href="/wiki/V-ATPase" title="V-ATPase">V-ATPase</a> generates a proton gradient at the expense of ATP, generating pH values of as low as 1.<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>The F<sub>1</sub> region also shows significant similarity to hexameric <a href="/wiki/DNA_helicase" class="mw-redirect" title="DNA helicase">DNA helicases</a> (especially the <a href="/wiki/Rho_factor" title="Rho factor">Rho factor</a>), and the entire enzyme region shows some similarity to <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span>-powered <a href="/wiki/T3SS" class="mw-redirect" title="T3SS">T3SS</a> or <a href="/wiki/Flagellum#Motor" title="Flagellum">flagellar motor</a> complexes.<sup id="cite_ref-InterPro_Synthase_18-1" class="reference"><a href="#cite_note-InterPro_Synthase-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><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><sup id="cite_ref-T3SS_21-0" class="reference"><a href="#cite_note-T3SS-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> The α<sub>3</sub>β<sub>3</sub> hexamer of the F<sub>1</sub> region shows significant structural similarity to hexameric DNA helicases; both form a ring with 3-fold rotational symmetry with a central pore. Both have roles dependent on the relative rotation of a macromolecule within the pore; the DNA helicases use the helical shape of DNA to drive their motion along the DNA molecule and to detect supercoiling, whereas the α<sub>3</sub>β<sub>3</sub> hexamer uses the conformational changes through the rotation of the γ subunit to drive an enzymatic reaction.<sup id="cite_ref-EctopicATP_22-0" class="reference"><a href="#cite_note-EctopicATP-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> motor of the F<sub>O</sub> particle shows great functional similarity to the <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> motors that drive flagella.<sup id="cite_ref-InterPro_Synthase_18-2" class="reference"><a href="#cite_note-InterPro_Synthase-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Both feature a ring of many small alpha-helical proteins that rotate relative to nearby stationary proteins, using a <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> potential gradient as an energy source. This link is tenuous, however, as the overall structure of flagellar motors is far more complex than that of the F<sub>O</sub> particle and the ring with about 30 rotating proteins is far larger than the 10, 11, or 14 helical proteins in the F<sub>O</sub> complex. More recent structural data do however show that the ring and the stalk are structurally similar to the F<sub>1</sub> particle.<sup id="cite_ref-T3SS_21-1" class="reference"><a href="#cite_note-T3SS-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/230px--ATP_synthesis_-_ATP_synthase_rotation.ogv.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="230" height="129" data-durationhint="19" data-mwtitle="ATP_synthesis_-_ATP_synthase_rotation.ogv" data-mwprovider="wikimediacommons" resource="/wiki/File:ATP_synthesis_-_ATP_synthase_rotation.ogv"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="854" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.720p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="720p.vp9.webm" data-width="1280" data-height="720" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.1080p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="1080p.vp9.webm" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv" type="video/ogg; codecs="theora"" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="256" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="426" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/1/18/ATP_synthesis_-_ATP_synthase_rotation.ogv/ATP_synthesis_-_ATP_synthase_rotation.ogv.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /></video></span><figcaption>Conformation changes of ATP synthase during synthesis</figcaption></figure> <p>The modular evolution theory for the origin of ATP synthase suggests that two subunits with independent function, a DNA helicase with ATPase activity and a <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> motor, were able to bind, and the rotation of the motor drove the ATPase activity of the helicase in reverse.<sup id="cite_ref-RotaryDNA_16-2" class="reference"><a href="#cite_note-RotaryDNA-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-EctopicATP_22-1" class="reference"><a href="#cite_note-EctopicATP-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> This complex then evolved greater efficiency and eventually developed into today's intricate ATP synthases. Alternatively, the DNA helicase/<span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> motor complex may have had <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> pump activity with the ATPase activity of the helicase driving the <span class="chemf nowrap">H<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span> motor in reverse.<sup id="cite_ref-RotaryDNA_16-3" class="reference"><a href="#cite_note-RotaryDNA-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> This may have evolved to carry out the reverse reaction and act as an ATP synthase.<sup id="cite_ref-LifeScience_IL_17-1" class="reference"><a href="#cite_note-LifeScience_IL-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cross1990_23-0" class="reference"><a href="#cite_note-Cross1990-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cross2004_24-0" class="reference"><a href="#cite_note-Cross2004-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Inhibitors">Inhibitors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=8" title="Edit section: Inhibitors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A variety of natural and synthetic inhibitors of ATP synthase have been discovered.<sup id="cite_ref-inhibitors_25-0" class="reference"><a href="#cite_note-inhibitors-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> These have been used to probe the structure and mechanism of ATP synthase. Some may be of therapeutic use. There are several classes of ATP synthase inhibitors, including peptide inhibitors, polyphenolic phytochemicals, polyketides, organotin compounds, polyenic α-pyrone derivatives, cationic inhibitors, substrate analogs, amino acid modifiers, and other miscellaneous chemicals.<sup id="cite_ref-inhibitors_25-1" class="reference"><a href="#cite_note-inhibitors-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Some of the most commonly used ATP synthase inhibitors are <a href="/wiki/Oligomycin" title="Oligomycin">oligomycin</a> and <a href="/wiki/N,N%27-Dicyclohexylcarbodiimide" title="N,N'-Dicyclohexylcarbodiimide">DCCD</a>. </p> <div class="mw-heading mw-heading2"><h2 id="In_different_organisms">In different organisms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=9" title="Edit section: In different organisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Bacteria">Bacteria</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=10" title="Edit section: Bacteria"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i><style data-mw-deduplicate="TemplateStyles:r1238216509">.mw-parser-output .vanchor>:target~.vanchor-text{background-color:#b1d2ff}@media screen{html.skin-theme-clientpref-night .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}</style><span class="vanchor"><span id="E._coli"></span><span class="vanchor-text">E. coli</span></span></i> ATP synthase is the simplest known form of ATP synthase, with 8 different subunit types.<sup id="cite_ref-ecoli_11-1" class="reference"><a href="#cite_note-ecoli-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Bacterial F-ATPases can occasionally operate in reverse, turning them into an ATPase.<sup id="cite_ref-pmid26671611_26-0" class="reference"><a href="#cite_note-pmid26671611-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Some bacteria have no F-ATPase, using an A/V-type ATPase bidirectionally.<sup id="cite_ref-pmid24878343_9-1" class="reference"><a href="#cite_note-pmid24878343-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Yeast">Yeast</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=11" title="Edit section: Yeast"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Yeast ATP synthase is one of the best-studied eukaryotic ATP synthases; and five F<sub>1</sub>, eight F<sub>O</sub> subunits, and seven associated proteins have been identified.<sup id="cite_ref-Velours_2000_7-1" class="reference"><a href="#cite_note-Velours_2000-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Most of these proteins have homologues in other eukaryotes.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Plant">Plant</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=12" title="Edit section: Plant"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In plants, ATP synthase is also present in <a href="/wiki/Chloroplasts" class="mw-redirect" title="Chloroplasts">chloroplasts</a> (CF<sub>1</sub>F<sub>O</sub>-ATP synthase). The enzyme is integrated into <a href="/wiki/Thylakoid" title="Thylakoid">thylakoid</a> membrane; the CF<sub>1</sub>-part sticks into <a href="/wiki/Stroma_(fluid)" title="Stroma (fluid)">stroma</a>, where dark reactions of photosynthesis (also called the light-independent reactions or the <a href="/wiki/Calvin_cycle" title="Calvin cycle">Calvin cycle</a>) and ATP synthesis take place. The overall structure and the catalytic mechanism of the chloroplast ATP synthase are almost the same as those of the bacterial enzyme. However, in chloroplasts, the <a href="/wiki/Electrochemical_potential" title="Electrochemical potential">proton motive force</a> is generated not by respiratory electron transport chain but by primary photosynthetic proteins. The synthase has a 40-aa insert in the gamma-subunit to inhibit wasteful activity when dark.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mammal">Mammal</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=13" title="Edit section: Mammal"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><span class="anchor" id="Bovine"></span>The ATP synthase isolated from bovine (<i>Bos taurus</i>) heart mitochondria is, in terms of biochemistry and structure, the best-characterized ATP synthase. Beef heart is used as a source for the enzyme because of the high concentration of mitochondria in cardiac muscle. Their genes have close homology to human ATP synthases.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p><span class="anchor" id="Human"></span>Human genes that encode components of ATP synthases: </p> <ul><li><i><a href="/wiki/ATP_synthase,_H%2B_transporting,_mitochondrial_F1_complex,_alpha_1" class="mw-redirect" title="ATP synthase, H+ transporting, mitochondrial F1 complex, alpha 1">ATP5A1</a></i></li> <li><i><a href="/wiki/ATP5B" title="ATP5B">ATP5B</a></i></li> <li><i><a href="/wiki/ATP5C1" title="ATP5C1">ATP5C1</a></i>, <i><a href="/wiki/ATP5D" title="ATP5D">ATP5D</a></i>, <i><a href="/wiki/ATP5E" title="ATP5E">ATP5E</a></i>, <i><a href="/wiki/ATP5F1" title="ATP5F1">ATP5F1</a></i>, <i><a href="/wiki/ATP5G1" title="ATP5G1">ATP5G1</a></i>, <i><a href="/wiki/ATP5G2" title="ATP5G2">ATP5G2</a></i>, <i><a href="/wiki/ATP5G3" title="ATP5G3">ATP5G3</a></i>, <i><a href="/wiki/ATP5H" title="ATP5H">ATP5H</a></i>, <i><a href="/wiki/ATP5I" title="ATP5I">ATP5I</a></i>, <i><a href="/wiki/ATP5J" class="mw-redirect" title="ATP5J">ATP5J</a></i>, <i><a href="/wiki/ATP5J2" title="ATP5J2">ATP5J2</a></i>, <i><a href="/wiki/ATP5L" title="ATP5L">ATP5L</a></i>, <i><a href="/wiki/ATP5O" class="mw-redirect" title="ATP5O">ATP5O</a></i></li> <li><i><a href="/wiki/MT-ATP6" title="MT-ATP6">MT-ATP6</a></i>, <i><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a></i></li></ul> <div class="mw-heading mw-heading3"><h3 id="Other_eukaryotes">Other eukaryotes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=14" title="Edit section: Other eukaryotes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Eukaryotes belonging to some divergent lineages have very special organizations of the ATP synthase. A <a href="/wiki/Euglenozoa" title="Euglenozoa">euglenozoa</a> ATP synthase forms a dimer with a boomerang-shaped F<sub>1</sub> head like other mitochondrial ATP synthases, but the F<sub>O</sub> subcomplex has many unique subunits. It uses <a href="/wiki/Cardiolipin" title="Cardiolipin">cardiolipin</a>. The inhibitory IF<sub>1</sub> also binds differently, in a way shared with <a href="/wiki/Trypanosomatida" title="Trypanosomatida">trypanosomatida</a>.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Archaea">Archaea</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=15" title="Edit section: Archaea"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Archaea do not generally have an F-ATPase. Instead, they synthesize ATP using the A-ATPase/synthase, a rotary machine structurally similar to the <a href="/wiki/V-ATPase" title="V-ATPase">V-ATPase</a> but mainly functioning as an ATP synthase.<sup id="cite_ref-pmid26671611_26-1" class="reference"><a href="#cite_note-pmid26671611-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Like the bacteria F-ATPase, it is believed to also function as an ATPase.<sup id="cite_ref-pmid24878343_9-2" class="reference"><a href="#cite_note-pmid24878343-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="LUCA_and_earlier">LUCA and earlier</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=16" title="Edit section: LUCA and earlier"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>F-ATPase gene linkage and gene order are widely conserved across ancient prokaryote lineages, implying that this system already existed at a date before the <a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">last universal common ancestor</a>, the LUCA.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<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=ATP_synthase&action=edit&section=17" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 25em;"> <ul><li><a href="/wiki/ATP10_protein" title="ATP10 protein">ATP10 protein</a> required for the assembly of the F<sub>O</sub> sector of the mitochondrial ATPase complex.</li> <li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Electron_transfer_chain" class="mw-redirect" title="Electron transfer chain">Electron transfer chain</a></li> <li><a href="/wiki/Flavoprotein" title="Flavoprotein">Flavoprotein</a></li> <li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">Oxidative phosphorylation</a></li> <li><a href="/wiki/P-ATPase" class="mw-redirect" title="P-ATPase">P-ATPase</a></li> <li><a href="/wiki/Proton_pump" title="Proton pump">Proton pump</a></li> <li><a href="/wiki/Rotating_locomotion_in_living_systems" title="Rotating locomotion in living systems">Rotating locomotion in living systems</a></li> <li><a href="/wiki/Transmembrane_ATPase" class="mw-redirect" title="Transmembrane ATPase">Transmembrane ATPase</a></li> <li><a href="/wiki/V-ATPase" title="V-ATPase">V-ATPase</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=18" 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 reflist-columns references-column-width" style="column-width: 33em;"> <ol class="references"> <li id="cite_note-Okuno_2011-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Okuno_2011_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation 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 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"Flagellar export apparatus and ATP synthetase: Homology evidenced by synteny predating the Last Universal Common Ancestor". <i>BioEssays</i>. <b>43</b> (7): e2100004. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fbies.202100004">10.1002/bies.202100004</a>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://hdl.handle.net/2292%2F55176">2292/55176</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/33998015">33998015</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:234747849">234747849</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BioEssays&rft.atitle=Flagellar+export+apparatus+and+ATP+synthetase%3A+Homology+evidenced+by+synteny+predating+the+Last+Universal+Common+Ancestor&rft.volume=43&rft.issue=7&rft.pages=e2100004&rft.date=2021-07&rft_id=info%3Ahdl%2F2292%2F55176&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A234747849%23id-name%3DS2CID&rft_id=info%3Apmid%2F33998015&rft_id=info%3Adoi%2F10.1002%2Fbies.202100004&rft.aulast=Matzke&rft.aufirst=NJ&rft.au=Lin%2C+A&rft.au=Stone%2C+M&rft.au=Baker%2C+MA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AATP+synthase" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=ATP_synthase&action=edit&section=19" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Nick Lane: <a rel="nofollow" class="external text" href="https://books.google.com/books?id=IfJYBQAAQBAJ&dq=nick+lane+the+vital+question+f0+f1+atp+synthase&pg=PT77"><i>The Vital Question: Energy, Evolution, and the Origins of Complex Life</i></a>, Ww Norton, 2015-07-20, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0393088816" title="Special:BookSources/978-0393088816">978-0393088816</a> (Link points to Figure 10 showing model of ATP synthase)</li></ul> <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=ATP_synthase&action=edit&section=20" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Boris A. Feniouk: <a rel="nofollow" class="external text" href="http://www.atpsynthase.info/">"ATP synthase — a splendid molecular machine"</a></li> <li>Well illustrated <a rel="nofollow" class="external text" href="http://www.life.uiuc.edu/crofts/bioph354/lect10.html">ATP synthase lecture</a> by Antony Crofts of the <a href="/wiki/University_of_Illinois_at_Urbana%E2%80%93Champaign" class="mw-redirect" title="University of Illinois at Urbana–Champaign">University of Illinois at Urbana–Champaign</a>.</li> <li><a rel="nofollow" class="external text" href="http://opm.phar.umich.edu/families.php?superfamily=5">Proton and Sodium translocating F-type, V-type and A-type ATPases in OPM database</a></li> <li><a rel="nofollow" class="external text" href="http://nobelprize.org/nobel_prizes/chemistry/laureates/1997/illpres/index.html">The Nobel Prize in Chemistry 1997</a> to Paul D. Boyer and John E. Walker for the enzymatic mechanism of synthesis of ATP; and to Jens C. Skou, for discovery of an ion-transporting enzyme, <span class="chemf nowrap">Na<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span>, <span class="chemf nowrap">K<span class="nowrap"><span style="display:inline-block;margin-bottom:-0.3em;vertical-align:0.8em;line-height:1em;font-size:80%;text-align:left"><sup style="font-size:inherit;line-height:inherit;vertical-align:baseline">+</sup><br /><sub style="font-size:inherit;line-height:inherit;vertical-align:baseline"></sub></span></span></span>-ATPase.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080422024412/http://multimedia.mcb.harvard.edu/media.html">Harvard Multimedia Production Site — Videos</a> – ATP synthesis animation</li> <li>David Goodsell: <a rel="nofollow" class="external text" href="http://www.rcsb.org/pdb/101/motm.do?momID=72">"ATP Synthase- Molecule of the Month"</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20150905064626/http://www.rcsb.org/pdb/101/motm.do?momID=72">Archived</a> 2015-09-05 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist 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.navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Membrane_transport_protein:_ion_pumps,_ATPases_/_ATP_synthase_(TC_3A2-3A3)" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:#e7dcc3"><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 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class="nv-edit"><a href="/wiki/Special:EditPage/Template:ATPase" title="Special:EditPage/Template:ATPase"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Membrane_transport_protein:_ion_pumps,_ATPases_/_ATP_synthase_(TC_3A2-3A3)" style="font-size:114%;margin:0 4em"><a href="/wiki/Membrane_transport_protein" title="Membrane transport protein">Membrane transport protein</a>: <a href="/wiki/Ion_transporter" title="Ion transporter">ion pumps</a>, <a href="/wiki/ATPase" title="ATPase">ATPases</a> / <a class="mw-selflink selflink">ATP synthase</a> (<a href="/wiki/Transporter_Classification_database#3.A._P-P-bond_hydrolysis-driven_transporters" class="mw-redirect" title="Transporter Classification database">TC 3A2-3A3</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink selflink">F-, V-, and A-type ATPase (3.A.2)</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%;background-color: AntiqueWhite"><a href="/wiki/F-ATPase" title="F-ATPase">H<sup>+</sup> (F-type)</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>H<sup>+</sup> transporting, mitochondrial: <a href="/wiki/ATP5A1" class="mw-redirect" title="ATP5A1">ATP5A1</a></li> <li><a href="/wiki/ATP5B" title="ATP5B">ATP5B</a></li> <li><a href="/wiki/ATP5C1" title="ATP5C1">ATP5C1</a></li> <li><a href="/w/index.php?title=ATP5C2&action=edit&redlink=1" class="new" title="ATP5C2 (page does not exist)">ATP5C2</a></li> <li><a href="/wiki/ATP5D" title="ATP5D">ATP5D</a></li> <li><a href="/wiki/ATP5E" title="ATP5E">ATP5E</a></li> <li><a href="/wiki/ATP5F1" title="ATP5F1">ATP5F1</a></li> <li><a href="/wiki/ATP5G1" title="ATP5G1">ATP5G1</a></li> <li><a href="/wiki/ATP5G2" title="ATP5G2">ATP5G2</a></li> <li><a href="/wiki/ATP5G3" title="ATP5G3">ATP5G3</a></li> <li><a href="/wiki/ATP5H" title="ATP5H">ATP5H</a></li> <li><a href="/wiki/ATP5I" title="ATP5I">ATP5I</a></li> <li><a href="/wiki/ATP5J" class="mw-redirect" title="ATP5J">ATP5J</a></li> <li><a href="/wiki/ATP5J2" title="ATP5J2">ATP5J2</a></li> <li><a href="/wiki/ATP5L" title="ATP5L">ATP5L</a></li> <li><a href="/w/index.php?title=ATP5L2&action=edit&redlink=1" class="new" title="ATP5L2 (page does not exist)">ATP5L2</a></li> <li><a href="/wiki/ATP5O" class="mw-redirect" title="ATP5O">ATP5O</a></li> <li><a href="/wiki/ATP5S" title="ATP5S">ATP5S</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/V-ATPase" title="V-ATPase">H<sup>+</sup> (V-type)</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/V-ATPase" title="V-ATPase">H<sup>+</sup> transporting, lysosomal</a>: <a href="/wiki/ATP6AP1" title="ATP6AP1">ATP6AP1</a></li> <li><a href="/wiki/ATP6AP2" class="mw-redirect" title="ATP6AP2">ATP6AP2</a></li> <li><a href="/wiki/ATP6V1A" title="ATP6V1A">ATP6V1A</a></li> <li><a href="/wiki/ATP6V1B1" title="ATP6V1B1">ATP6V1B1</a></li> <li><a href="/wiki/ATP6V1B2" title="ATP6V1B2">ATP6V1B2</a></li> <li><a href="/wiki/ATP6V1C1" title="ATP6V1C1">ATP6V1C1</a></li> <li><a href="/wiki/ATP6V1C2" title="ATP6V1C2">ATP6V1C2</a></li> <li><a href="/wiki/ATP6V1D" title="ATP6V1D">ATP6V1D</a></li> <li><a href="/wiki/ATP6V1E1" title="ATP6V1E1">ATP6V1E1</a></li> <li><a href="/wiki/ATP6V1E2" title="ATP6V1E2">ATP6V1E2</a></li> <li><a href="/wiki/ATP6V1F" title="ATP6V1F">ATP6V1F</a></li> <li><a href="/wiki/ATP6V1G1" title="ATP6V1G1">ATP6V1G1</a></li> <li><a href="/wiki/ATP6V1G2" title="ATP6V1G2">ATP6V1G2</a></li> <li><a href="/wiki/ATP6V1G3" title="ATP6V1G3">ATP6V1G3</a></li> <li><a href="/wiki/ATP6V1H" title="ATP6V1H">ATP6V1H</a></li> <li><a href="/wiki/ATP6V0A1" class="mw-redirect" title="ATP6V0A1">ATP6V0A1</a></li> <li><a href="/wiki/ATP6V0A2" title="ATP6V0A2">ATP6V0A2</a></li> <li><a href="/wiki/ATP6V0A4" title="ATP6V0A4">ATP6V0A4</a></li> <li><a href="/wiki/ATP6V0B" title="ATP6V0B">ATP6V0B</a></li> <li><a href="/wiki/ATP6V0C" title="ATP6V0C">ATP6V0C</a></li> <li><a href="/wiki/ATP6V0D1" title="ATP6V0D1">ATP6V0D1</a></li> <li><a href="/wiki/ATP6V0D2" title="ATP6V0D2">ATP6V0D2</a></li> <li><a href="/w/index.php?title=ATP6V0E&action=edit&redlink=1" class="new" title="ATP6V0E (page does not exist)">ATP6V0E</a></li> <li><a href="/wiki/ATP6V0E1" title="ATP6V0E1">ATP6V0E1</a></li></ul> <ul><li><a href="/wiki/TCIRG1" title="TCIRG1">TCIRG1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/ATPase#Classification" title="ATPase">A-ATPase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">found in Archea</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/P-type_ATPase" title="P-type ATPase">P-type ATPase</a> (3.A.3)</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>3.A.3.1.1: <a href="/wiki/Na%2B/K%2B-ATPase" class="mw-redirect" title="Na+/K+-ATPase">Na<sup>+</sup>/K<sup>+</sup> transporting</a>: <a href="/wiki/ATP1A1" class="mw-redirect" title="ATP1A1">ATP1A1</a></li> <li><a href="/wiki/ATP1A2" title="ATP1A2">ATP1A2</a></li> <li><a href="/wiki/ATP1A3" title="ATP1A3">ATP1A3</a></li> <li><a href="/wiki/ATP1A4" title="ATP1A4">ATP1A4</a></li> <li><a href="/wiki/ATP1B1" title="ATP1B1">ATP1B1</a></li> <li><a href="/w/index.php?title=ATP1B2&action=edit&redlink=1" class="new" title="ATP1B2 (page does not exist)">ATP1B2</a></li> <li><a href="/wiki/ATP1B3" title="ATP1B3">ATP1B3</a></li> <li><a href="/w/index.php?title=ATP1B4&action=edit&redlink=1" class="new" title="ATP1B4 (page does not exist)">ATP1B4</a></li> <li><a href="/w/index.php?title=ATP1G1&action=edit&redlink=1" class="new" title="ATP1G1 (page does not exist)">ATP1G1</a></li></ul> <ul><li>3.A.3.1.2: <a href="/wiki/Hydrogen_potassium_ATPase" title="Hydrogen potassium ATPase">H<sup>+</sup>/K<sup>+</sup></a></li> <li><a href="/wiki/Hydrogen_potassium_ATPase" title="Hydrogen potassium ATPase">H<sup>+</sup>/K<sup>+</sup> exchanging</a>: <a href="/w/index.php?title=ATP4A&action=edit&redlink=1" class="new" title="ATP4A (page does not exist)">ATP4A</a></li> <li><a href="/w/index.php?title=ATP4B&action=edit&redlink=1" class="new" title="ATP4B (page does not exist)">ATP4B</a></li></ul> <ul><li>3.A.3.1.4: H<sup>+</sup>/K<sup>+</sup> transporting, nongastric: <a href="/wiki/ATP12A" title="ATP12A">ATP12A</a></li></ul> <ul><li>3.A.3.2: <a href="/wiki/Calcium_ATPase" title="Calcium ATPase">Ca<sup>2+</sup></a> (<a href="/wiki/SERCA" title="SERCA">SERCA</a>, <a href="/wiki/Plasma_membrane_Ca%C2%B2%E2%81%BA_ATPase" class="mw-redirect" title="Plasma membrane Ca²⁺ ATPase">PMCA</a>, <a href="/wiki/Secretory_Pathway_Ca%C2%B2%E2%81%BA_ATPase" title="Secretory Pathway Ca²⁺ ATPase">SPCA</a>) / <a href="/wiki/Calcium_ATPase" title="Calcium ATPase">Ca<sup>2+</sup> transporting</a>: <a href="/wiki/ATP2A1" title="ATP2A1">ATP2A1</a></li> <li><a href="/wiki/ATP2A2" title="ATP2A2">ATP2A2</a></li> <li><a href="/wiki/ATP2A3" title="ATP2A3">ATP2A3</a></li> <li><a href="/wiki/ATP2B1" title="ATP2B1">ATP2B1</a></li> <li><a href="/wiki/ATP2B2" title="ATP2B2">ATP2B2</a></li> <li><a href="/wiki/ATP2B3" title="ATP2B3">ATP2B3</a></li> <li><a href="/wiki/ATP2B4" title="ATP2B4">ATP2B4</a></li> <li><a href="/wiki/ATP2C1" title="ATP2C1">ATP2C1</a></li></ul> <ul><li>3.A.3.5: Cu<sup>2+</sup> transporting: <a href="/wiki/ATP7A" title="ATP7A">ATP7A</a></li> <li><a href="/wiki/Wilson_disease_protein" title="Wilson disease protein">ATP7B</a></li></ul> <ul><li>3.A.3.8.8: <a href="/wiki/Flippase" title="Flippase">flippase</a>: <a href="/w/index.php?title=ATP8A2&action=edit&redlink=1" class="new" title="ATP8A2 (page does not exist)">ATP8A2</a></li></ul> <ul><li>Other/ungrouped:</li></ul> <ul><li><a href="/wiki/Na%2B/K%2B-ATPase" class="mw-redirect" title="Na+/K+-ATPase">Na<sup>+</sup>/K<sup>+</sup></a> – <a href="/wiki/Proton_ATPase" title="Proton ATPase">H<sup>+</sup></a></li></ul> <ul><li><a href="/wiki/Mg%2B%2B_transporting" class="mw-redirect" title="Mg++ transporting">Mg<sup>2+</sup> transporting</a>: <a href="/wiki/ATP3" class="mw-redirect" title="ATP3">ATP3</a></li> <li>Class I, type 8: <a href="/w/index.php?title=ATP8A1&action=edit&redlink=1" class="new" title="ATP8A1 (page does not exist)">ATP8A1</a></li> <li><a href="/wiki/ATP8B1" class="mw-redirect" title="ATP8B1">ATP8B1</a></li> <li><a href="/w/index.php?title=ATP8B2&action=edit&redlink=1" class="new" title="ATP8B2 (page does not exist)">ATP8B2</a></li> <li><a href="/wiki/ATP8B3" title="ATP8B3">ATP8B3</a></li> <li><a href="/w/index.php?title=ATP8B4&action=edit&redlink=1" class="new" title="ATP8B4 (page does not exist)">ATP8B4</a></li> <li>Class II, type 9: <a href="/wiki/ATP9A" title="ATP9A">ATP9A</a></li> <li><a href="/w/index.php?title=ATP9B&action=edit&redlink=1" class="new" title="ATP9B (page does not exist)">ATP9B</a></li> <li>Class V, type 10: <a href="/wiki/ATP10A" class="mw-redirect" title="ATP10A">ATP10A</a></li> <li><a href="/w/index.php?title=ATP10B&action=edit&redlink=1" class="new" title="ATP10B (page does not exist)">ATP10B</a></li> <li><a href="/w/index.php?title=ATP10D&action=edit&redlink=1" class="new" title="ATP10D (page does not exist)">ATP10D</a></li> <li>Class VI, type 11: <a href="/w/index.php?title=ATP11A&action=edit&redlink=1" class="new" title="ATP11A (page does not exist)">ATP11A</a></li> <li><a href="/wiki/ATP11B" title="ATP11B">ATP11B</a></li> <li><a href="/wiki/ATP11C" title="ATP11C">ATP11C</a></li> <li>type 13: <a href="/w/index.php?title=ATP13A1&action=edit&redlink=1" class="new" title="ATP13A1 (page does not exist)">ATP13A1</a></li> <li><a href="/wiki/ATP13A2" title="ATP13A2">ATP13A2</a></li> <li><a href="/wiki/ATP13A3" title="ATP13A3">ATP13A3</a></li> <li><a href="/w/index.php?title=ATP13A4&action=edit&redlink=1" class="new" title="ATP13A4 (page does not exist)">ATP13A4</a></li> <li><a href="/w/index.php?title=ATP13A5&action=edit&redlink=1" class="new" title="ATP13A5 (page does not exist)">ATP13A5</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background-color: AntiqueWhite"><div><i>see also <a href="/wiki/Template:ATPase_disorders" title="Template:ATPase disorders">ATPase disorders</a></i></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="Mitochondrial_proteins" 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:Mitochondrial_proteins" title="Template:Mitochondrial proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Mitochondrial_proteins" title="Template talk:Mitochondrial proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Mitochondrial_proteins" title="Special:EditPage/Template:Mitochondrial proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Mitochondrial_proteins" style="font-size:114%;margin:0 4em"><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrial</a> proteins</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Outer_mitochondrial_membrane" class="mw-redirect" title="Outer mitochondrial membrane">Outer membrane</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%"><span class="nobold"><i><a href="/wiki/Fatty_acid_degradation" title="Fatty acid degradation">fatty acid degradation</a></i></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><a href="/wiki/Carnitine_palmitoyltransferase_I" title="Carnitine palmitoyltransferase I">Carnitine palmitoyltransferase I</a></li> <li><a href="/wiki/Long-chain-fatty-acid%E2%80%94CoA_ligase" title="Long-chain-fatty-acid—CoA ligase">Long-chain-fatty-acid—CoA ligase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> metabolism</i></span></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/Kynureninase" title="Kynureninase">Kynureninase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Monoamine_neurotransmitter" title="Monoamine neurotransmitter">monoamine neurotransmitter</a><br /> metabolism</i></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><a href="/wiki/Monoamine_oxidase" title="Monoamine oxidase">Monoamine oxidase</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/Mitochondrial_intermembrane_space" class="mw-redirect" title="Mitochondrial intermembrane space">Intermembrane space</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/Adenylate_kinase" title="Adenylate kinase">Adenylate kinase</a></li> <li><a href="/wiki/Creatine_kinase" title="Creatine kinase">Creatine kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">Inner membrane</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%"><span class="nobold"><i><a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a></i></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><a href="/wiki/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Coenzyme Q – cytochrome c reductase</a></li> <li><a href="/wiki/Cytochrome_c" title="Cytochrome c">Cytochrome c</a></li> <li><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a></li> <li><a href="/wiki/Succinate_dehydrogenase" title="Succinate dehydrogenase">Succinate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Pyrimidine_metabolism" title="Pyrimidine metabolism">pyrimidine metabolism</a></i></span></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/Dihydroorotate_dehydrogenase" title="Dihydroorotate dehydrogenase">Dihydroorotate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Mitochondrial_shuttle" title="Mitochondrial shuttle">mitochondrial shuttle</a></i></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><a href="/wiki/Malate-aspartate_shuttle" class="mw-redirect" title="Malate-aspartate shuttle">Malate-aspartate shuttle</a></li> <li><a href="/wiki/Glycerol_phosphate_shuttle" title="Glycerol phosphate shuttle">Glycerol phosphate shuttle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Steroidogenesis" class="mw-redirect" title="Steroidogenesis">steroidogenesis</a></i></span></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/Cholesterol_side-chain_cleavage_enzyme" title="Cholesterol side-chain cleavage enzyme">Cholesterol side-chain cleavage enzyme</a></li> <li><a href="/wiki/Steroid_11-beta-hydroxylase" class="mw-redirect" title="Steroid 11-beta-hydroxylase">Steroid 11-beta-hydroxylase</a></li> <li><a href="/wiki/Aldosterone_synthase" title="Aldosterone synthase">Aldosterone synthase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>other</i></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><a href="/wiki/Glutamate_aspartate_transporter" class="mw-redirect" title="Glutamate aspartate transporter">Glutamate aspartate transporter</a></li> <li><a href="/wiki/Glycerol-3-phosphate_dehydrogenase" title="Glycerol-3-phosphate dehydrogenase">Glycerol-3-phosphate dehydrogenase</a></li> <li><a class="mw-selflink selflink">ATP synthase</a></li> <li><a href="/wiki/Carnitine_palmitoyltransferase_II" title="Carnitine palmitoyltransferase II">Carnitine palmitoyltransferase II</a></li> <li><a href="/wiki/Uncoupling_protein" title="Uncoupling protein">Uncoupling protein</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/Mitochondrial_matrix" title="Mitochondrial matrix">Matrix</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%"><span class="nobold"><i><a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</a></i></span></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/Citrate_synthase" title="Citrate synthase">Citrate synthase</a></li> <li><a href="/wiki/Aconitase" title="Aconitase">Aconitase</a></li> <li><a href="/wiki/Isocitrate_dehydrogenase" title="Isocitrate dehydrogenase">Isocitrate dehydrogenase</a></li> <li><a href="/wiki/Oxoglutarate_dehydrogenase_complex" title="Oxoglutarate dehydrogenase complex">Oxoglutarate dehydrogenase complex</a></li> <li><a href="/wiki/Succinyl_coenzyme_A_synthetase" title="Succinyl coenzyme A synthetase">Succinyl coenzyme A synthetase</a></li> <li><a href="/wiki/Fumarase" title="Fumarase">Fumarase</a></li> <li><a href="/wiki/Malate_dehydrogenase" title="Malate dehydrogenase">Malate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Anaplerotic_reactions" title="Anaplerotic reactions">anaplerotic reactions</a></i></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><a href="/wiki/Aspartate_transaminase" title="Aspartate transaminase">Aspartate transaminase</a></li> <li><a href="/wiki/Glutamate_dehydrogenase" title="Glutamate dehydrogenase">Glutamate dehydrogenase</a></li> <li><a href="/wiki/Pyruvate_dehydrogenase_complex" title="Pyruvate dehydrogenase complex">Pyruvate dehydrogenase complex</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Urea_cycle" title="Urea cycle">urea cycle</a></i></span></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/Carbamoyl_phosphate_synthetase_I" title="Carbamoyl phosphate synthetase I">Carbamoyl phosphate synthetase I</a></li> <li><a href="/wiki/Ornithine_transcarbamylase" title="Ornithine transcarbamylase">Ornithine transcarbamylase</a></li> <li><a href="/wiki/N-Acetylglutamate_synthase" title="N-Acetylglutamate synthase">N-Acetylglutamate synthase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>alcohol metabolism</i></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><a href="/wiki/ALDH2" title="ALDH2">ALDH2</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/PMPCB" title="PMPCB">PMPCB</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/to be sorted</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><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Frataxin" title="Frataxin">Frataxin</a></li> <li><a href="/wiki/Mitochondrial_membrane_transport_protein" title="Mitochondrial membrane transport protein">Mitochondrial membrane transport protein</a> <ul><li><a href="/wiki/Mitochondrial_permeability_transition_pore" title="Mitochondrial permeability transition pore">Mitochondrial permeability transition pore</a></li> <li><a href="/wiki/Mitochondrial_carrier" title="Mitochondrial carrier">Mitochondrial carrier</a></li></ul></li> <li><a href="/wiki/Translocator_protein" title="Translocator protein">Translocator protein</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/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial DNA</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%"><span class="nobold"><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">Complex I</a></span></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/MT-ND1" title="MT-ND1">MT-ND1</a></li> <li><a href="/wiki/MT-ND2" title="MT-ND2">MT-ND2</a></li> <li><a href="/wiki/MT-ND3" title="MT-ND3">MT-ND3</a></li> <li><a href="/wiki/MT-ND4" title="MT-ND4">MT-ND4</a></li> <li><a href="/wiki/MT-ND4L" title="MT-ND4L">MT-ND4L</a></li> <li><a href="/wiki/MT-ND5" title="MT-ND5">MT-ND5</a></li> <li><a href="/wiki/MT-ND6" title="MT-ND6">MT-ND6</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Complex III</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><a href="/wiki/MT-CYB" title="MT-CYB">MT-CYB</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Complex IV</a></span></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/Cytochrome_c_oxidase_subunit_I" title="Cytochrome c oxidase subunit I">MT-CO1</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_II" class="mw-redirect" title="Cytochrome c oxidase subunit II">MT-CO2</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_III" title="Cytochrome c oxidase subunit III">MT-CO3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a class="mw-selflink selflink">ATP synthase</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><a href="/wiki/MT-ATP6" title="MT-ATP6">MT-ATP6</a></li> <li><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Transfer_RNA" title="Transfer RNA">tRNA</a></span></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/MT-TA" title="MT-TA">MT-TA</a></li> <li><a href="/wiki/MT-TC" title="MT-TC">MT-TC</a></li> <li><a href="/wiki/MT-TD" title="MT-TD">MT-TD</a></li> <li><a href="/wiki/MT-TE" title="MT-TE">MT-TE</a></li> <li><a href="/wiki/MT-TF" title="MT-TF">MT-TF</a></li> <li><a href="/wiki/MT-TG" title="MT-TG">MT-TG</a></li> <li><a href="/wiki/MT-TH" title="MT-TH">MT-TH</a></li> <li><a href="/wiki/MT-TI" title="MT-TI">MT-TI</a></li> <li><a href="/wiki/MT-TK" title="MT-TK">MT-TK</a></li> <li><a href="/wiki/MT-TL1" title="MT-TL1">MT-TL1</a></li> <li><a href="/wiki/MT-TL2" title="MT-TL2">MT-TL2</a></li> <li><a href="/wiki/MT-TM" title="MT-TM">MT-TM</a></li> <li><a href="/wiki/MT-TN" title="MT-TN">MT-TN</a></li> <li><a href="/wiki/MT-TP" title="MT-TP">MT-TP</a></li> <li><a href="/wiki/MT-TQ" title="MT-TQ">MT-TQ</a></li> <li><a href="/wiki/MT-TR" title="MT-TR">MT-TR</a></li> <li><a href="/wiki/MT-TS1" title="MT-TS1">MT-TS1</a></li> <li><a href="/wiki/MT-TS2" title="MT-TS2">MT-TS2</a></li> <li><a href="/wiki/MT-TT" title="MT-TT">MT-TT</a></li> <li><a href="/wiki/MT-TV_(mitochondrial)" title="MT-TV (mitochondrial)">MT-TV</a></li> <li><a href="/wiki/MT-TW" title="MT-TW">MT-TW</a></li> <li><a href="/wiki/MT-TY" title="MT-TY">MT-TY</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div><i>see also <a href="/wiki/Template:Mitochondrial_diseases" title="Template:Mitochondrial diseases">mitochondrial diseases</a></i></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="Hydrolases:_acid_anhydride_hydrolases_(EC_3.6)" 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:Acid_anhydride_hydrolases" title="Template:Acid anhydride hydrolases"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Acid_anhydride_hydrolases" title="Template talk:Acid anhydride hydrolases"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Acid_anhydride_hydrolases" title="Special:EditPage/Template:Acid anhydride hydrolases"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Hydrolases:_acid_anhydride_hydrolases_(EC_3.6)" style="font-size:114%;margin:0 4em"><a href="/wiki/Hydrolase" title="Hydrolase">Hydrolases</a>: <a href="/wiki/Acid_anhydride_hydrolases" title="Acid anhydride hydrolases">acid anhydride hydrolases</a> (<a href="/wiki/Enzyme_Commission_number" title="Enzyme Commission number">EC</a> 3.6)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#3.6.1" title="List of EC numbers (EC 3)">3.6.1</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/Pyrophosphatase" title="Pyrophosphatase">Pyrophosphatase</a> <ul><li><a href="/wiki/Inorganic_pyrophosphatase" title="Inorganic pyrophosphatase">Inorganic</a></li> <li><a href="/wiki/Thiamine_pyrophosphatase" class="mw-redirect" title="Thiamine pyrophosphatase">Thiamine</a></li></ul></li> <li><a href="/wiki/Apyrase" title="Apyrase">Apyrase</a></li> <li><a href="/wiki/Thiamine-triphosphatase" title="Thiamine-triphosphatase">Thiamine-triphosphatase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#3.6.2" title="List of EC numbers (EC 3)">3.6.2</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/Adenylylsulfatase" title="Adenylylsulfatase">Adenylylsulfatase</a></li> <li><a href="/wiki/Phosphoadenylylsulfatase" title="Phosphoadenylylsulfatase">Phosphoadenylylsulfatase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#3.6.3" title="List of EC numbers (EC 3)">3.6.3</a>-<a href="/wiki/List_of_EC_numbers_(EC_3)#3.6.4" title="List of EC numbers (EC 3)">4</a>: <a href="/wiki/ATPase" title="ATPase">ATPase</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%">3.6.3</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%">Cu++ (3.6.3.4)</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/ATP7A" title="ATP7A">Menkes/ATP7A</a></li> <li><a href="/wiki/Wilson_disease_protein" title="Wilson disease protein">Wilson/ATP7B</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Calcium_ATPase" title="Calcium ATPase">Ca+</a> (3.6.3.8)</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/SERCA" title="SERCA">SERCA</a> <ul><li><a href="/wiki/ATP2A1" title="ATP2A1">ATP2A1</a></li> <li><a href="/wiki/ATP2A2" title="ATP2A2">ATP2A2</a></li> <li><a href="/wiki/ATP2A3" title="ATP2A3">ATP2A3</a></li></ul></li> <li><a href="/wiki/Plasma_membrane_Ca2%2B_ATPase" title="Plasma membrane Ca2+ ATPase">Plasma membrane</a> <ul><li><a href="/wiki/ATP2B1" title="ATP2B1">ATP2B1</a></li> <li><a href="/wiki/ATP2B2" title="ATP2B2">ATP2B2</a></li> <li><a href="/wiki/ATP2B3" title="ATP2B3">ATP2B3</a></li> <li><a href="/wiki/ATP2B4" title="ATP2B4">ATP2B4</a></li></ul></li> <li><a href="/wiki/Secretory_Pathway_Ca%C2%B2%E2%81%BA_ATPase" title="Secretory Pathway Ca²⁺ ATPase">SPCA</a> <ul><li><a href="/wiki/ATP2C1" title="ATP2C1">ATP2C1</a></li> <li><a href="/w/index.php?title=ATP2C2&action=edit&redlink=1" class="new" title="ATP2C2 (page does not exist)">ATP2C2</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Na%2B/K%2B-ATPase" class="mw-redirect" title="Na+/K+-ATPase">Na+/K+</a> (3.6.3.9)</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/ATPase,_Na%2B/K%2B_transporting,_alpha_1" title="ATPase, Na+/K+ transporting, alpha 1">ATP1A1</a></li> <li><a href="/wiki/ATP1A2" title="ATP1A2">ATP1A2</a></li> <li><a href="/wiki/ATP1A3" title="ATP1A3">ATP1A3</a></li> <li><a href="/wiki/ATP1A4" title="ATP1A4">ATP1A4</a></li> <li><a href="/wiki/ATP1B1" title="ATP1B1">ATP1B1</a></li> <li><a href="/w/index.php?title=ATP1B2&action=edit&redlink=1" class="new" title="ATP1B2 (page does not exist)">ATP1B2</a></li> <li><a href="/wiki/ATP1B3" title="ATP1B3">ATP1B3</a></li> <li><a href="/w/index.php?title=ATP1B4&action=edit&redlink=1" class="new" title="ATP1B4 (page does not exist)">ATP1B4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hydrogen_potassium_ATPase" title="Hydrogen potassium ATPase">H+/K+</a> (3.6.3.10)</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="/w/index.php?title=ATP4A&action=edit&redlink=1" class="new" title="ATP4A (page does not exist)">ATP4A</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other <a href="/wiki/P-type_ATPase" title="P-type ATPase">P-type ATPase</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/ATP8B1" class="mw-redirect" title="ATP8B1">ATP8B1</a></li> <li><a href="/wiki/ATP10A" class="mw-redirect" title="ATP10A">ATP10A</a></li> <li><a href="/wiki/ATP11B" title="ATP11B">ATP11B</a></li> <li><a href="/wiki/ATP12A" title="ATP12A">ATP12A</a></li> <li><a href="/wiki/ATP13A2" title="ATP13A2">ATP13A2</a></li> <li><a href="/wiki/ATP13A3" title="ATP13A3">ATP13A3</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">3.6.4</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/Dynein" title="Dynein">Dynein</a></li> <li><a href="/wiki/Kinesin" title="Kinesin">Kinesin</a></li> <li><a href="/wiki/Myosin" title="Myosin">Myosin</a></li> <li><a href="/wiki/Katanin" title="Katanin">Katanin</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/List_of_EC_numbers_(EC_3)#3.6.5" title="List of EC numbers (EC 3)">3.6.5</a>: <a href="/wiki/GTPase" title="GTPase">GTPase</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%">3.6.5.1: <a href="/wiki/Heterotrimeric_G_protein" title="Heterotrimeric G protein">Heterotrimeric G protein</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/Gs_alpha_subunit" title="Gs alpha subunit">G<sub>αs</sub></a> <ul><li><a href="/wiki/Olfactory_receptor" title="Olfactory receptor">G<sub>olf</sub></a></li></ul></li> <li><a href="/wiki/Gi_alpha_subunit" title="Gi alpha subunit">G<sub>αi</sub></a> <ul><li><a href="/wiki/GNAI1" title="GNAI1">GNAI1</a></li> <li><a href="/wiki/GNAI2" title="GNAI2">GNAI2</a></li> <li><a href="/wiki/GNAI3" title="GNAI3">GNAI3</a></li> <li><a href="/wiki/Transducin" title="Transducin">Transducin</a> <ul><li><a href="/wiki/GNAT1" title="GNAT1">GNAT1</a></li> <li><a href="/wiki/GNAT2" title="GNAT2">GNAT2</a></li></ul></li> <li><a href="/wiki/Gustducin" title="Gustducin">Gustducin</a> <ul><li><a href="/wiki/GNAT3" title="GNAT3">GNAT3</a></li></ul></li></ul></li> <li><a href="/wiki/Gq_alpha_subunit" title="Gq alpha subunit">G<sub>αq/11</sub></a> <ul><li><a href="/wiki/GNAQ" title="GNAQ">GNAQ</a></li> <li><a href="/wiki/GNA11" title="GNA11">GNA11</a></li></ul></li> <li><a href="/wiki/G12/G13_alpha_subunits" title="G12/G13 alpha subunits">G<sub>α12/13</sub></a> <ul><li><a href="/wiki/GNA12" title="GNA12">GNA12</a></li> <li><a href="/wiki/GNA13" title="GNA13">GNA13</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">3.6.5.2: <a href="/wiki/Small_GTPase" title="Small GTPase">Small GTPase</a> > <a href="/wiki/Ras_superfamily" title="Ras superfamily">Ras superfamily</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/Rho_family_of_GTPases" title="Rho family of GTPases">Rho family of GTPases</a>: <i>Cdc42</i> <ul><li><a href="/wiki/CDC42" title="CDC42">CDC42</a></li> <li><a href="/wiki/TC10_protein" class="mw-redirect" title="TC10 protein">TC10</a></li> <li><a href="/wiki/TCL_(GTPase)" title="TCL (GTPase)">TCL</a></li></ul></li> <li><i>RhoUV</i> <ul><li><a href="/wiki/Wrch1" title="Wrch1">RhoU</a></li> <li><a href="/wiki/Chp_(GTPase)" title="Chp (GTPase)">RhoV</a></li></ul></li> <li><a href="/wiki/Rac_(GTPase)" title="Rac (GTPase)">Rac</a> <ul><li><a href="/wiki/RAC1" title="RAC1">Rac1</a></li> <li><a href="/wiki/RAC2" title="RAC2">2</a></li> <li><a href="/wiki/Rac3" class="mw-redirect" title="Rac3">3</a></li> <li><a href="/wiki/RhoG" title="RhoG">RhoG</a></li></ul></li> <li><i>RhoBTB</i> <ul><li><a href="/wiki/RHOBTB1" title="RHOBTB1">1</a></li> <li><a href="/wiki/RHOBTB2" title="RHOBTB2">2</a></li></ul></li> <li><a href="/wiki/RhoH" title="RhoH">RhoH</a></li> <li><a href="/wiki/Rho_family_of_GTPases" title="Rho family of GTPases">Rho</a> <ul><li><a href="/wiki/RHOA" class="mw-redirect" title="RHOA">A</a></li> <li><a href="/wiki/RHOB" title="RHOB">B</a></li> <li><a href="/wiki/RhoC" title="RhoC">C</a></li></ul></li> <li><a href="/wiki/Rnd_(GTPase)" title="Rnd (GTPase)">Rnd</a> <ul><li><a href="/wiki/Rnd1" title="Rnd1">1</a></li> <li><a href="/wiki/Rnd2" title="Rnd2">2</a></li> <li><a href="/wiki/Rnd3" title="Rnd3">3</a></li></ul></li> <li><i>RhoDF</i> <ul><li><a href="/wiki/Rif_(GTPase)" title="Rif (GTPase)">RhoF</a></li> <li><a href="/wiki/RhoD" title="RhoD">RhoD</a></li></ul></li></ul> <ul><li>other: <a href="/wiki/Ras_subfamily" class="mw-redirect" title="Ras subfamily">Ras</a> <ul><li><a href="/wiki/HRAS" title="HRAS">HRAS</a></li> <li><a href="/wiki/KRAS" title="KRAS">KRAS</a></li> <li><a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a></li></ul></li> <li><a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rab</a> <ul><li><a href="/wiki/RAB23" title="RAB23">RAB23</a></li> <li><a href="/wiki/RAB27" title="RAB27">RAB27</a></li></ul></li> <li><a href="/wiki/ADP_ribosylation_factor" title="ADP ribosylation factor">Arf</a> <ul><li><a href="/wiki/ARF6" title="ARF6">ARF6</a></li> <li><a href="/wiki/SAR1B" class="mw-redirect" title="SAR1B">SAR1B</a></li> <li><a href="/wiki/ARL13B" title="ARL13B">ARL13B</a></li> <li><a href="/wiki/ARL6" title="ARL6">ARL6</a></li></ul></li> <li><a href="/wiki/Ran_(gene)" class="mw-redirect" title="Ran (gene)">Ran</a></li> <li><a href="/wiki/RHEB" title="RHEB">Rheb</a></li> <li><a href="/wiki/Rap_GTP-binding_protein" title="Rap GTP-binding protein">Rap</a></li> <li><a href="/wiki/RRAD_(gene)" title="RRAD (gene)">RGK</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">3.6.5.3: <a href="/wiki/Protein-synthesizing_GTPase" title="Protein-synthesizing GTPase">Protein-synthesizing GTPase</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/Prokaryotic_elongation_factors" class="mw-redirect" title="Prokaryotic elongation factors">Prokaryotic</a> <ul><li><a href="/wiki/Prokaryotic_initiation_factor-2" class="mw-redirect" title="Prokaryotic initiation factor-2">IF-2</a></li> <li><a href="/wiki/EF-Tu" title="EF-Tu">EF-Tu</a></li> <li><a href="/wiki/EF-G" title="EF-G">EF-G</a></li></ul></li> <li><a href="/wiki/Eukaryotic_elongation_factors" class="mw-redirect" title="Eukaryotic elongation factors">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">3.6.5.5-6: <a href="/wiki/Molecular_motor" title="Molecular motor">Polymerization motors</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/Dynamin_superfamily" title="Dynamin superfamily">dynamin superfamily</a> <ul><li><a href="/wiki/Dynamin" title="Dynamin">Dynamin</a></li> <li><a href="/wiki/Guanylate-binding_protein" title="Guanylate-binding protein">Guanylate-binding protein</a></li> <li><a href="/w/index.php?title=Mitofusin-1&action=edit&redlink=1" class="new" title="Mitofusin-1 (page does not exist)">Mitofusin-1</a></li> <li><a href="/wiki/MX1" title="MX1">MX1</a> and <a href="/wiki/MX2" title="MX2">MX2</a></li> <li><a href="/wiki/Dynamin-like_120_kDa_protein" title="Dynamin-like 120 kDa protein">OPA1</a></li></ul></li> <li><a href="/wiki/Tubulin" title="Tubulin">Tubulin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Enzymes" 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:Enzymes" title="Template:Enzymes"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Enzymes" title="Template talk:Enzymes"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Enzymes" title="Special:EditPage/Template:Enzymes"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Enzymes" style="font-size:114%;margin:0 4em"><a href="/wiki/Enzyme" title="Enzyme">Enzymes</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Activity</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/Active_site" title="Active site">Active site</a></li> <li><a href="/wiki/Binding_site" title="Binding site">Binding site</a></li> <li><a href="/wiki/Catalytic_triad" title="Catalytic triad">Catalytic triad</a></li> <li><a href="/wiki/Oxyanion_hole" title="Oxyanion hole">Oxyanion hole</a></li> <li><a href="/wiki/Enzyme_promiscuity" title="Enzyme promiscuity">Enzyme promiscuity</a></li> <li><a href="/wiki/Diffusion-limited_enzyme" title="Diffusion-limited enzyme">Diffusion-limited enzyme</a></li> <li><a href="/wiki/Cofactor_(biochemistry)" title="Cofactor (biochemistry)">Cofactor</a></li> <li><a href="/wiki/Enzyme_catalysis" title="Enzyme catalysis">Enzyme catalysis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Regulation</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/Allosteric_regulation" title="Allosteric regulation">Allosteric regulation</a></li> <li><a href="/wiki/Cooperativity" title="Cooperativity">Cooperativity</a></li> <li><a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">Enzyme inhibitor</a></li> <li><a href="/wiki/Enzyme_activator" title="Enzyme activator">Enzyme activator</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Classification</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/Enzyme_Commission_number" title="Enzyme Commission number">EC number</a></li> <li><a href="/wiki/Protein_superfamily" title="Protein superfamily">Enzyme superfamily</a></li> <li><a href="/wiki/Protein_family" title="Protein family">Enzyme family</a></li> <li><a href="/wiki/List_of_enzymes" title="List of enzymes">List of enzymes</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Kinetics</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/Enzyme_kinetics" title="Enzyme kinetics">Enzyme kinetics</a></li> <li><a href="/wiki/Eadie%E2%80%93Hofstee_diagram" title="Eadie–Hofstee diagram">Eadie–Hofstee diagram</a></li> <li><a href="/wiki/Hanes%E2%80%93Woolf_plot" title="Hanes–Woolf plot">Hanes–Woolf plot</a></li> <li><a href="/wiki/Lineweaver%E2%80%93Burk_plot" title="Lineweaver–Burk plot">Lineweaver–Burk plot</a></li> <li><a href="/wiki/Michaelis%E2%80%93Menten_kinetics" title="Michaelis–Menten kinetics">Michaelis–Menten kinetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</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><b>EC1 <a href="/wiki/Oxidoreductase" title="Oxidoreductase">Oxidoreductases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_1)" title="List of EC numbers (EC 1)">list</a>)</li> <li><b>EC2 <a href="/wiki/Transferase" title="Transferase">Transferases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_2)" title="List of EC numbers (EC 2)">list</a>)</li> <li><b>EC3 <a href="/wiki/Hydrolase" 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