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<span>Mechanism</span> </div> </a> <ul id="toc-Mechanism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-G_domain_GTPases" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#G_domain_GTPases"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>G domain GTPases</span> </div> </a> <button aria-controls="toc-G_domain_GTPases-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 G domain GTPases subsection</span> </button> <ul id="toc-G_domain_GTPases-sublist" class="vector-toc-list"> <li id="toc-TRAFAC_class" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#TRAFAC_class"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>TRAFAC class</span> </div> </a> <ul id="toc-TRAFAC_class-sublist" class="vector-toc-list"> <li id="toc-Translation_factor_superfamily" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Translation_factor_superfamily"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.1</span> <span>Translation factor superfamily</span> </div> </a> <ul id="toc-Translation_factor_superfamily-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ras-like_superfamily" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Ras-like_superfamily"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.2</span> <span>Ras-like superfamily</span> </div> </a> <ul id="toc-Ras-like_superfamily-sublist" class="vector-toc-list"> <li id="toc-Heterotrimeric_G_proteins" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Heterotrimeric_G_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.2.1</span> <span>Heterotrimeric G proteins</span> </div> </a> <ul id="toc-Heterotrimeric_G_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Small_GTPases" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Small_GTPases"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.2.2</span> <span>Small GTPases</span> </div> </a> <ul id="toc-Small_GTPases-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Myosin-kinesin_superfamily" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Myosin-kinesin_superfamily"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.3</span> <span>Myosin-kinesin superfamily</span> </div> </a> <ul id="toc-Myosin-kinesin_superfamily-sublist" class="vector-toc-list"> <li id="toc-Large_GTPases" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Large_GTPases"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.3.1</span> <span>Large GTPases</span> </div> </a> <ul id="toc-Large_GTPases-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-SIMIBI_class" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#SIMIBI_class"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>SIMIBI class</span> </div> </a> <ul id="toc-SIMIBI_class-sublist" class="vector-toc-list"> <li id="toc-Translocation_factors" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Translocation_factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Translocation factors</span> </div> </a> <ul id="toc-Translocation_factors-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Other_GTPases" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Other_GTPases"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Other GTPases</span> </div> </a> <ul id="toc-Other_GTPases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-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">7</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet 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Available in 16 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-16" 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">16 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%A8%D8%B1%D9%88%D8%AA%D9%8A%D9%86_%D8%A7%D9%84GTPase" title="بروتين الGTPase – Arabic" lang="ar" hreflang="ar" data-title="بروتين الGTPase" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/GTPaza" title="GTPaza – Bosnian" lang="bs" hreflang="bs" data-title="GTPaza" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/GTP%C3%A1za" title="GTPáza – Czech" lang="cs" hreflang="cs" data-title="GTPáza" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/GTPasa" title="GTPasa – Spanish" lang="es" hreflang="es" data-title="GTPasa" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AC%DB%8C%E2%80%8C%D8%AA%DB%8C%E2%80%8C%D9%BE%DB%8C%E2%80%8C%D8%A2%D8%B2" title="جی‌تی‌پی‌آز – Persian" lang="fa" hreflang="fa" data-title="جی‌تی‌پی‌آز" 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/GTPase" title="GTPase – French" lang="fr" hreflang="fr" data-title="GTPase" 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/GTPase" title="GTPase – Galician" lang="gl" hreflang="gl" data-title="GTPase" 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/GTP%EA%B0%80%EC%88%98%EB%B6%84%ED%95%B4%ED%9A%A8%EC%86%8C" title="GTP가수분해효소 – Korean" lang="ko" hreflang="ko" data-title="GTP가수분해효소" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/GTPasi" title="GTPasi – Italian" lang="it" hreflang="it" data-title="GTPasi" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/GTPase" title="GTPase – Dutch" lang="nl" hreflang="nl" data-title="GTPase" 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/GTP%E3%82%A2%E3%83%BC%E3%82%BC" title="GTPアーゼ – Japanese" lang="ja" hreflang="ja" data-title="GTPアーゼ" 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-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/GTPase" title="GTPase – Portuguese" lang="pt" hreflang="pt" data-title="GTPase" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%93%D0%A2%D0%A4%D0%B0%D0%B7%D0%B0" title="ГТФаза – Russian" lang="ru" hreflang="ru" data-title="ГТФаза" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/GTPaze" title="GTPaze – Serbian" lang="sr" hreflang="sr" data-title="GTPaze" 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/GTPaze" title="GTPaze – Serbo-Croatian" lang="sh" hreflang="sh" data-title="GTPaze" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Guanosiinitrifosfataasit" title="Guanosiinitrifosfataasit – Finnish" lang="fi" hreflang="fi" data-title="Guanosiinitrifosfataasit" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li> </ul> <div 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a large family of <a href="/wiki/Hydrolase" title="Hydrolase">hydrolase</a> <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that bind to the <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">guanosine triphosphate (GTP)</a> and <a href="/wiki/Hydrolysis" title="Hydrolysis">hydrolyze</a> it to <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">guanosine diphosphate (GDP)</a>.<sup id="cite_ref-pmid8462668_1-0" class="reference"><a href="#cite_note-pmid8462668-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The GTP binding and hydrolysis takes place in the highly <a href="/wiki/Conserved_sequence" title="Conserved sequence">conserved</a> <a href="/wiki/P-loop" class="mw-redirect" title="P-loop">P-loop</a> "G domain", a <a href="/wiki/Domain_(protein)" class="mw-redirect" title="Domain (protein)">protein domain</a> common to many GTPases.<sup id="cite_ref-pmid8462668_1-1" class="reference"><a href="#cite_note-pmid8462668-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Functions">Functions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=1" title="Edit section: Functions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>GTPases function as molecular switches or timers in many fundamental cellular processes.<sup id="cite_ref-pmid3113327_2-0" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>Examples of these roles include: </p> <ul><li><a href="/wiki/Signal_transduction" title="Signal transduction">Signal transduction</a> in response to activation of cell surface receptors, including <a href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">transmembrane receptors</a> such as those mediating <a href="/wiki/Taste" title="Taste">taste</a>, <a href="/wiki/Olfaction" class="mw-redirect" title="Olfaction">smell</a> and <a href="/wiki/Visual_perception" title="Visual perception">vision</a>.<sup id="cite_ref-pmid3113327_2-1" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">Protein biosynthesis</a> (a.k.a. <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>) at the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>.</li> <li>Regulation of cell <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a>, <a href="/wiki/Cell_growth" title="Cell growth">proliferation</a>, <a href="/wiki/Cell_division" title="Cell division">division</a> and <a href="/wiki/Cell_migration" title="Cell migration">movement</a>.</li> <li><a href="/wiki/Protein_targeting#Protein_translocation" title="Protein targeting">Translocation</a> of <a href="/wiki/Protein" title="Protein">proteins</a> through <a href="/wiki/Cell_membrane" title="Cell membrane">membranes</a>.</li> <li>Transport of <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a> within the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>, and vesicle-mediated secretion and uptake, through GTPase control of vesicle coat assembly.</li></ul> <p>GTPases are active when bound to GTP and inactive when bound to GDP.<sup id="cite_ref-pmid3113327_2-2" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-0" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In the generalized receptor-transducer-effector signaling model of <a href="/wiki/Martin_Rodbell" title="Martin Rodbell">Martin Rodbell</a>, signaling GTPases act as transducers to regulate the activity of effector proteins.<sup id="cite_ref-pmid7579038_3-1" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> This inactive-active switch is due to conformational changes in the protein distinguishing these two forms, particularly of the "switch" regions that in the active state are able to make protein-protein contacts with partner proteins that alter the function of these effectors.<sup id="cite_ref-pmid8462668_1-2" class="reference"><a href="#cite_note-pmid8462668-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=2" title="Edit section: Mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Hydrolysis of GTP bound to an (active) G domain-GTPase leads to deactivation of the signaling/timer function of the enzyme.<sup id="cite_ref-pmid3113327_2-3" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-2" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The hydrolysis of the third (γ) <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> of GTP to create <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">guanosine diphosphate</a> (GDP) and P<sub>i</sub>, <a href="/wiki/Inorganic_phosphate" class="mw-redirect" title="Inorganic phosphate">inorganic phosphate</a>, occurs by the S<sub>N</sub>2 mechanism (see <a href="/wiki/Nucleophilic_substitution" title="Nucleophilic substitution">nucleophilic substitution</a>) via a pentacoordinate transition state and is dependent on the presence of a <a href="/wiki/Magnesium" title="Magnesium">magnesium</a> <a href="/wiki/Ion" title="Ion">ion</a> Mg<sup>2+</sup>.<sup id="cite_ref-pmid3113327_2-4" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-3" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>GTPase activity serves as the shutoff mechanism for the signaling roles of GTPases by returning the active, GTP-bound protein to the inactive, GDP-bound state.<sup id="cite_ref-pmid3113327_2-5" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-4" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Most "GTPases" have functional GTPase activity, allowing them to remain active (that is, bound to GTP) only for a short time before deactivating themselves by converting bound GTP to bound GDP.<sup id="cite_ref-pmid3113327_2-6" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-5" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> However, many GTPases also use accessory proteins named <a href="/wiki/GTPase-activating_proteins" class="mw-redirect" title="GTPase-activating proteins">GTPase-activating proteins</a> or GAPs to accelerate their GTPase activity. This further limits the active lifetime of signaling GTPases.<sup id="cite_ref-pmid9430654_4-0" class="reference"><a href="#cite_note-pmid9430654-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Some GTPases have little to no intrinsic GTPase activity, and are entirely dependent on GAP proteins for deactivation (such as the <a href="/wiki/ADP-ribosylation_factor" class="mw-redirect" title="ADP-ribosylation factor">ADP-ribosylation factor</a> or ARF family of small GTP-binding proteins that are involved in vesicle-mediated transport within cells).<sup id="cite_ref-pmid3086320_5-0" class="reference"><a href="#cite_note-pmid3086320-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>To become activated, GTPases must bind to GTP. Since mechanisms to convert bound GDP directly into GTP are unknown, the inactive GTPases are induced to release bound GDP by the action of distinct regulatory proteins called <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">guanine nucleotide exchange factors</a> or GEFs.<sup id="cite_ref-pmid3113327_2-7" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-6" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The nucleotide-free GTPase protein quickly rebinds GTP, which is in far excess in healthy cells over GDP, allowing the GTPase to enter the active conformation state and promote its effects on the cell.<sup id="cite_ref-pmid3113327_2-8" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-7" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> For many GTPases, activation of GEFs is the primary control mechanism in the stimulation of the GTPase signaling functions, although GAPs also play an important role. For heterotrimeric G proteins and many small GTP-binding proteins, GEF activity is stimulated by cell surface receptors in response to signals outside the cell (for heterotrimeric G proteins, the <a href="/wiki/G_protein-coupled_receptors" class="mw-redirect" title="G protein-coupled receptors">G protein-coupled receptors</a> are themselves GEFs, while for receptor-activated small GTPases their GEFs are distinct from cell surface receptors). </p><p>Some GTPases also bind to accessory proteins called <a href="/wiki/Guanosine_nucleotide_dissociation_inhibitor" title="Guanosine nucleotide dissociation inhibitor">guanine nucleotide dissociation inhibitors</a> or GDIs that stabilize the inactive, GDP-bound state.<sup id="cite_ref-pmid9588168_6-0" class="reference"><a href="#cite_note-pmid9588168-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The amount of active GTPase can be changed in several ways: </p> <ol><li>Acceleration of GDP dissociation by GEFs speeds up the accumulation of active GTPase.</li> <li>Inhibition of GDP dissociation by guanine nucleotide dissociation inhibitors (GDIs) slows down accumulation of active GTPase.</li> <li>Acceleration of GTP hydrolysis by GAPs reduces the amount of active GTPase.</li> <li>Artificial <i>GTP analogues</i> like <i>GTP-γ-S</i>, <i>β,γ-methylene-GTP</i>, and <i>β,γ-imino-GTP</i> that cannot be hydrolyzed can lock the GTPase in its active state.</li> <li>Mutations (such as those that reduce the intrinsic GTP hydrolysis rate) can lock the GTPase in the active state, and such mutations in the small GTPase Ras are particularly common in some forms of cancer.<sup id="cite_ref-pmid31255772_7-0" class="reference"><a href="#cite_note-pmid31255772-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup></li></ol> <div class="mw-heading mw-heading2"><h2 id="G_domain_GTPases">G domain GTPases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=3" title="Edit section: G domain GTPases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In most GTPases, the specificity for the base <a href="/wiki/Guanine" title="Guanine">guanine</a> versus other nucleotides is imparted by the base-recognition motif, which has the consensus sequence [N/T]KXD. The following classification is based on shared features; some examples have mutations in the base-recognition motif that shift their substrate specificity, most commonly to ATP.<sup id="cite_ref-pmid11916378_8-0" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="TRAFAC_class">TRAFAC class</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=4" title="Edit section: TRAFAC class"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The TRAFAC class of G domain proteins is named after the prototypical member, the translation factor G proteins. They play roles in translation, signal transduction, and cell motility.<sup id="cite_ref-pmid11916378_8-1" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Translation_factor_superfamily">Translation factor superfamily<span class="anchor" id="Translation_factor_family"></span></h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=5" title="Edit section: Translation factor superfamily"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/EF-G#Evolution" title="EF-G">EF-G § Evolution</a>, and <a href="/wiki/EF-Tu#Evolution" title="EF-Tu">EF-Tu § Evolution</a></div> <p>Multiple classical <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> factor family GTPases play important roles in <a href="/wiki/Transcription_(genetics)#initiation" class="mw-redirect" title="Transcription (genetics)">initiation</a>, <a href="/wiki/Eukaryotic_translation#Elongation" title="Eukaryotic translation">elongation</a> and termination of <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein biosynthesis</a>. Sharing a similar mode of <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> binding due to the β-EI domain following the GTPase, the most well-known members of the family are <a href="/wiki/EEF-1" title="EEF-1">EF-1A</a>/<a href="/wiki/EF-Tu" title="EF-Tu">EF-Tu</a>, <a href="/wiki/EEF-2" class="mw-redirect" title="EEF-2">EF-2</a>/<a href="/wiki/EF-G" title="EF-G">EF-G</a>,<sup id="cite_ref-pmid6113539_9-0" class="reference"><a href="#cite_note-pmid6113539-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> and class 2 <a href="/wiki/Release_factor" title="Release factor">release factors</a>. Other members include <a href="/wiki/EF-4" title="EF-4">EF-4</a> (LepA), <a href="/w/index.php?title=BipA&action=edit&redlink=1" class="new" title="BipA (page does not exist)">BipA</a> (TypA),<sup id="cite_ref-pmid29235176_10-0" class="reference"><a href="#cite_note-pmid29235176-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> <i>SelB</i> (bacterial selenocysteinyl-tRNA EF-Tu paralog), <i>Tet</i> (<a href="/wiki/Tetracycline" title="Tetracycline">tetracycline</a> resistance by ribosomal protection),<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/HBS1L" class="mw-redirect" title="HBS1L">HBS1L</a> (eukaryotic <a href="/wiki/Ribosomal_pause" title="Ribosomal pause">ribosome rescue</a> protein similar to release factors). </p><p>The superfamily also includes the Bms1 family from yeast.<sup id="cite_ref-pmid11916378_8-2" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Ras-like_superfamily">Ras-like superfamily</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=6" title="Edit section: Ras-like superfamily"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading5"><h5 id="Heterotrimeric_G_proteins">Heterotrimeric G proteins</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=7" title="Edit section: Heterotrimeric G proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Heterotrimeric_G_protein" title="Heterotrimeric G protein">Heterotrimeric G protein</a></div> <p><a href="/wiki/Heterotrimeric_G_protein" title="Heterotrimeric G protein">Heterotrimeric G protein</a> complexes are composed of three distinct protein subunits named <i><a href="/wiki/G_alpha_subunit" title="G alpha subunit">alpha</a></i> (α), <i>beta</i> (β) and <i>gamma</i> (γ) <a href="/wiki/Protein_subunit" title="Protein subunit">subunits</a>.<sup id="cite_ref-pmid10819326_12-0" class="reference"><a href="#cite_note-pmid10819326-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The alpha subunits contain the GTP binding/GTPase domain flanked by long regulatory regions, while the beta and gamma subunits form a stable dimeric complex referred to as the <a href="/wiki/G_beta-gamma_complex" title="G beta-gamma complex">beta-gamma complex</a>.<sup id="cite_ref-pmid9131251_13-0" class="reference"><a href="#cite_note-pmid9131251-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> When activated, a heterotrimeric G protein dissociates into activated, GTP-bound alpha subunit and separate beta-gamma subunit, each of which can perform distinct signaling roles.<sup id="cite_ref-pmid3113327_2-9" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-8" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The α and γ subunit are modified by <a href="/wiki/Lipid_anchor" class="mw-redirect" title="Lipid anchor">lipid anchors</a> to increase their association with the inner leaflet of the plasma membrane.<sup id="cite_ref-pmid11313912_14-0" class="reference"><a href="#cite_note-pmid11313912-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>Heterotrimeric G proteins act as the transducers of <a href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">G protein-coupled receptors</a>, coupling receptor activation to downstream signaling effectors and <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">second messengers</a>.<sup id="cite_ref-pmid3113327_2-10" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-9" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid12209124_15-0" class="reference"><a href="#cite_note-pmid12209124-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In unstimulated cells, heterotrimeric G proteins are assembled as the GDP bound, inactive trimer (G<sub>α</sub>-GDP-G<sub>βγ</sub> complex).<sup id="cite_ref-pmid3113327_2-11" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-10" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Upon receptor activation, the activated receptor intracellular domain acts as GEF to release GDP from the G protein complex and to promote binding of GTP in its place.<sup id="cite_ref-pmid3113327_2-12" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-11" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The GTP-bound complex undergoes an activating conformation shift that dissociates it from the receptor and also breaks the complex into its component G protein alpha and beta-gamma subunit components.<sup id="cite_ref-pmid3113327_2-13" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-12" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> While these activated G protein subunits are now free to activate their effectors, the active receptor is likewise free to activate additional G proteins – this allows catalytic activation and amplification where one receptor may activate many G proteins.<sup id="cite_ref-pmid3113327_2-14" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-13" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>G protein signaling is terminated by hydrolysis of bound GTP to bound GDP.<sup id="cite_ref-pmid3113327_2-15" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-14" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> This can occur through the intrinsic GTPase activity of the α subunit, or be accelerated by separate regulatory proteins that act as <a href="/wiki/GTPase-activating_protein" title="GTPase-activating protein">GTPase-activating proteins</a> (GAPs), such as members of the <a href="/wiki/Regulator_of_G_protein_signaling" title="Regulator of G protein signaling">Regulator of G protein signaling</a> (RGS) family).<sup id="cite_ref-pmid9430654_4-1" class="reference"><a href="#cite_note-pmid9430654-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The speed of the hydrolysis reaction works as an internal clock limiting the length of the signal. Once G<sub>α</sub> is returned to being GDP bound, the two parts of the heterotrimer re-associate to the original, inactive state.<sup id="cite_ref-pmid3113327_2-16" class="reference"><a href="#cite_note-pmid3113327-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7579038_3-15" class="reference"><a href="#cite_note-pmid7579038-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The heterotrimeric G proteins can be classified by <a href="/wiki/Sequence_homology" title="Sequence homology">sequence homology</a> of the α unit and by their functional targets into four families: G<sub>s</sub> family, G<sub>i</sub> family, G<sub>q</sub> family and G<sub>12</sub> family.<sup id="cite_ref-pmid10819326_12-1" class="reference"><a href="#cite_note-pmid10819326-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Each of these G<sub>α</sub> protein families contains multiple members, such that the mammals have 16 distinct <sub>α</sub>-subunit genes.<sup id="cite_ref-pmid10819326_12-2" class="reference"><a href="#cite_note-pmid10819326-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The G<sub>β</sub> and G<sub>γ</sub> are likewise composed of many members, increasing heterotrimer structural and functional diversity.<sup id="cite_ref-pmid10819326_12-3" class="reference"><a href="#cite_note-pmid10819326-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Among the target molecules of the specific G proteins are the second messenger-generating enzymes <a href="/wiki/Adenylyl_cyclase" title="Adenylyl cyclase">adenylyl cyclase</a> and <a href="/wiki/Phospholipase_C" title="Phospholipase C">phospholipase C</a>, as well as various <a href="/wiki/Ion_channel" title="Ion channel">ion channels</a>.<sup id="cite_ref-pmid12040175_16-0" class="reference"><a href="#cite_note-pmid12040175-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading5"><h5 id="Small_GTPases">Small GTPases</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=8" title="Edit section: Small GTPases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Small_GTPase" title="Small GTPase">Small GTPase</a></div> <p><a href="/wiki/Small_GTPase" title="Small GTPase">Small GTPases</a> function as monomers and have a molecular weight of about 21 kilodaltons that consists primarily of the GTPase domain.<sup id="cite_ref-pmid11152757_17-0" class="reference"><a href="#cite_note-pmid11152757-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> They are also called small or monomeric guanine nucleotide-binding regulatory proteins, small or monomeric GTP-binding proteins, or small or monomeric G-proteins, and because they have significant homology with the first-identified such protein, named <a href="/wiki/Ras_(protein)" class="mw-redirect" title="Ras (protein)">Ras</a>, they are also referred to as <a href="/wiki/Ras_superfamily" title="Ras superfamily">Ras superfamily</a> GTPases. Small GTPases generally serve as molecular switches and signal transducers for a wide variety of cellular signaling events, often involving membranes, vesicles or cytoskeleton.<sup id="cite_ref-pmid2116664_18-0" class="reference"><a href="#cite_note-pmid2116664-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11152757_17-1" class="reference"><a href="#cite_note-pmid11152757-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> According to their primary amino acid sequences and biochemical properties, the many Ras superfamily small GTPases are further divided into five subfamilies with distinct functions: <a href="/wiki/Ras_(protein)" class="mw-redirect" title="Ras (protein)">Ras</a>, <a href="/wiki/Rho_family_of_GTPases" title="Rho family of GTPases">Rho</a> ("Ras-homology"), <a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rab</a>, <a href="/wiki/ADP_ribosylation_factor" title="ADP ribosylation factor">Arf</a> and <a href="/wiki/Ran_protein" class="mw-redirect" title="Ran protein">Ran</a>.<sup id="cite_ref-pmid11152757_17-2" class="reference"><a href="#cite_note-pmid11152757-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> While many small GTPases are activated by their GEFs in response to intracellular signals emanating from cell surface receptors (particularly <a href="/wiki/Growth_factor_receptor" title="Growth factor receptor">growth factor receptors</a>), regulatory GEFs for many other small GTPases are activated in response to intrinsic cell signals, not cell surface (external) signals. </p> <div class="mw-heading mw-heading4"><h4 id="Myosin-kinesin_superfamily">Myosin-kinesin superfamily</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=9" title="Edit section: Myosin-kinesin superfamily"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This class is defined by loss of two beta-strands and additional N-terminal strands. Both namesakes of this superfamily, <a href="/wiki/Myosin" title="Myosin">myosin</a> and <a href="/wiki/Kinesin" title="Kinesin">kinesin</a>, have shifted to use ATP.<sup id="cite_ref-pmid11916378_8-3" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading5"><h5 id="Large_GTPases">Large GTPases</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=10" title="Edit section: Large GTPases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>See <a href="/wiki/Dynamin" title="Dynamin">dynamin</a> as a prototype for large monomeric GTPases. </p> <div class="mw-heading mw-heading3"><h3 id="SIMIBI_class">SIMIBI class</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=11" title="Edit section: SIMIBI class"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Much of the SIMIBI class of GTPases is activated by dimerization.<sup id="cite_ref-pmid11916378_8-4" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Named after the signal recognition particle (SRP), MinD, and BioD, the class is involved in protein localization, chromosome partitioning, and membrane transport. Several members of this class, including MinD and Get3, has shifted in substrate specificity to become ATPases.<sup id="cite_ref-pmid27658684_19-0" class="reference"><a href="#cite_note-pmid27658684-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Translocation_factors">Translocation factors</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=12" title="Edit section: Translocation factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Missing_information plainlinks metadata ambox ambox-content" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Wiki_letter_w.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/44px-Wiki_letter_w.svg.png" decoding="async" width="44" height="44" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/66px-Wiki_letter_w.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/88px-Wiki_letter_w.svg.png 2x" data-file-width="44" data-file-height="44" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>is missing information</b> about FlhF, which is not involved in the SRP at all but has a similar structure to Ffh and FtsY.<span class="hide-when-compact"> Please expand the section to include this information. Further details may exist on the <a href="/wiki/Talk:GTPase" title="Talk:GTPase">talk page</a>.</span> <span class="date-container"><i>(<span class="date">October 2021</span>)</i></span></div></td></tr></tbody></table> <p>For a discussion of <a href="/wiki/Protein_translocation" class="mw-redirect" title="Protein translocation">Translocation</a> factors and the role of GTP, see <a href="/wiki/Signal_recognition_particle" title="Signal recognition particle">signal recognition particle</a> (SRP). </p> <div class="mw-heading mw-heading2"><h2 id="Other_GTPases">Other GTPases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=13" title="Edit section: Other GTPases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While <a href="/wiki/Tubulin" title="Tubulin">tubulin</a> and related structural proteins also bind and hydrolyze GTP as part of their function to form intracellular tubules, these proteins utilize a distinct <a href="/wiki/Tubulin_domain" title="Tubulin domain">tubulin domain</a> that is unrelated to the G domain used by signaling GTPases.<sup id="cite_ref-pmid9628483_20-0" class="reference"><a href="#cite_note-pmid9628483-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>There are also GTP-hydrolyzing proteins that use a <a href="/wiki/P-loop" class="mw-redirect" title="P-loop">P-loop</a> from a superclass other than the G-domain-containing one. Examples include the <a href="/wiki/NACHT_domain" title="NACHT domain">NACHT</a> proteins of its own superclass and McrB protein of the <a href="/wiki/AAA%2B" class="mw-redirect" title="AAA+">AAA+</a> superclass.<sup id="cite_ref-pmid11916378_8-5" class="reference"><a href="#cite_note-pmid11916378-8"><span class="cite-bracket">[</span>8<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=GTPase&action=edit&section=14" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/G_protein-coupled_receptors" class="mw-redirect" title="G protein-coupled receptors">G protein-coupled receptors</a></li> <li><a href="/wiki/Growth_factor_receptor" title="Growth factor receptor">Growth factor receptor</a></li> <li><a href="/wiki/Septins" class="mw-redirect" title="Septins">Septins</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=GTPase&action=edit&section=15" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-pmid8462668-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid8462668_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid8462668_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-pmid8462668_1-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFStoutenSanderWittinghoferValencia1993" class="citation journal cs1">Stouten, PF; Sander, C; Wittinghofer, A; Valencia, A (1993). "How does the switch II region of G-domains work?". <i>FEBS Letters</i>. <b>320</b> (1): <span class="nowrap">1–</span>6. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F0014-5793%2893%2981644-f">10.1016/0014-5793(93)81644-f</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/8462668">8462668</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=FEBS+Letters&rft.atitle=How+does+the+switch+II+region+of+G-domains+work%3F&rft.volume=320&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1-%3C%2Fspan%3E6&rft.date=1993&rft_id=info%3Adoi%2F10.1016%2F0014-5793%2893%2981644-f&rft_id=info%3Apmid%2F8462668&rft.aulast=Stouten&rft.aufirst=PF&rft.au=Sander%2C+C&rft.au=Wittinghofer%2C+A&rft.au=Valencia%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGTPase" class="Z3988"></span></span> </li> <li id="cite_note-pmid3113327-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid3113327_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-12"><sup><i><b>m</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-13"><sup><i><b>n</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-14"><sup><i><b>o</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-15"><sup><i><b>p</b></i></sup></a> <a href="#cite_ref-pmid3113327_2-16"><sup><i><b>q</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGilman1987" class="citation journal cs1">Gilman, AG (1987). 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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 class="mw-selflink selflink">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="Enzymes56" 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 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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" title="Hydrolase">Hydrolases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_3)" title="List of EC numbers (EC 3)">list</a>)</li> <li><b>EC4 <a href="/wiki/Lyase" title="Lyase">Lyases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_4)" title="List of EC numbers (EC 4)">list</a>)</li> <li><b>EC5 <a href="/wiki/Isomerase" title="Isomerase">Isomerases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_5)" title="List of EC numbers (EC 5)">list</a>)</li> <li><b>EC6 <a href="/wiki/Ligase" title="Ligase">Ligases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_6)" title="List of EC numbers (EC 6)">list</a>)</li> <li><b>EC7 <a href="/wiki/Translocase" title="Translocase">Translocases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_7)" title="List of EC numbers (EC 7)">list</a>)</li></ul> </div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;justify-content:center;align-items:baseline}.mw-parser-output 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