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</ul> </li> <li id="toc-Membrane_attachment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Membrane_attachment"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Membrane attachment</span> </div> </a> <ul id="toc-Membrane_attachment-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Members" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Members"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Members</span> </div> </a> <ul id="toc-Members-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ras_in_cancer" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Ras_in_cancer"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Ras in cancer</span> </div> </a> <button aria-controls="toc-Ras_in_cancer-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 Ras in cancer subsection</span> </button> <ul id="toc-Ras_in_cancer-sublist" class="vector-toc-list"> <li id="toc-Inappropriate_activation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inappropriate_activation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Inappropriate activation</span> </div> </a> <ul id="toc-Inappropriate_activation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Constitutively_active_Ras" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Constitutively_active_Ras"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Constitutively active Ras</span> </div> </a> <ul id="toc-Constitutively_active_Ras-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ras-targeted_cancer_treatments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ras-targeted_cancer_treatments"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Ras-targeted cancer treatments</span> </div> </a> <ul id="toc-Ras-targeted_cancer_treatments-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-In_other_species" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#In_other_species"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>In other species</span> </div> </a> <button aria-controls="toc-In_other_species-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 other species subsection</span> </button> <ul id="toc-In_other_species-sublist" class="vector-toc-list"> <li id="toc-Xenopus_laevis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Xenopus_laevis"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span><i>Xenopus laevis</i></span> </div> </a> <ul id="toc-Xenopus_laevis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Drosophila_melanogaster" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Drosophila_melanogaster"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span><i>Drosophila melanogaster</i></span> </div> </a> <ul id="toc-Drosophila_melanogaster-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Aplysia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Aplysia"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span><i>Aplysia</i></span> </div> </a> <ul id="toc-Aplysia-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Caenorhabditis_elegans" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Caenorhabditis_elegans"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span><i>Caenorhabditis elegans</i></span> </div> </a> <ul id="toc-Caenorhabditis_elegans-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dictyostelium_discoideum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dictyostelium_discoideum"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.5</span> <span><i>Dictyostelium discoideum</i></span> </div> </a> <ul id="toc-Dictyostelium_discoideum-sublist" class="vector-toc-list"> </ul> </li> </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">7</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">8</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">9</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 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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 15 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-15" 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">15 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%B9%D8%A7%D8%A6%D9%84%D8%A9_%D8%B1%D8%A7%D8%B3" title="عائلة راس – Arabic" lang="ar" hreflang="ar" data-title="عائلة راس" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Ras_(protein)" title="Ras (protein) – Czech" lang="cs" hreflang="cs" data-title="Ras (protein)" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/G-Protein_Ras" title="G-Protein Ras – German" lang="de" hreflang="de" data-title="G-Protein Ras" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Ras" title="Ras – Spanish" lang="es" hreflang="es" data-title="Ras" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Prot%C3%A9ine_Ras" title="Protéine Ras – French" lang="fr" hreflang="fr" data-title="Protéine Ras" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Ras_(proteina)" title="Ras (proteina) – Italian" lang="it" hreflang="it" data-title="Ras (proteina)" 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/Ras" title="Ras – Hebrew" lang="he" hreflang="he" data-title="Ras" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Ras_(eiwit)" title="Ras (eiwit) – Dutch" lang="nl" hreflang="nl" data-title="Ras (eiwit)" 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/Ras%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA" title="Rasタンパク質 – Japanese" lang="ja" hreflang="ja" data-title="Rasタンパク質" 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/Prote%C3%ADnas_RAS" title="Proteínas RAS – Portuguese" lang="pt" hreflang="pt" data-title="Proteínas RAS" 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/Ras" title="Ras – Russian" lang="ru" hreflang="ru" data-title="Ras" 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/Ras_potfamilija" title="Ras potfamilija – Serbian" lang="sr" hreflang="sr" data-title="Ras potfamilija" 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/Ras_potfamilija" title="Ras potfamilija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Ras potfamilija" 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-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Ras_(protein)" title="Ras (protein) – Turkish" lang="tr" hreflang="tr" data-title="Ras (protein)" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B1%D8%A7%D8%B3_(%D8%B3%D8%A7%D9%84%D9%85%D8%A7%D8%AA%DB%8C_%D8%AD%DB%8C%D8%A7%D8%AA%DB%8C%D8%A7%D8%AA)" title="راس (سالماتی حیاتیات) – Urdu" lang="ur" hreflang="ur" data-title="راس (سالماتی حیاتیات)" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q424396#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div 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.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">This article is about p21/Ras protein. For the p21/waf1 protein, see <a href="/wiki/P21" title="P21">p21</a>.</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><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Hras_surface_colored_by_conservation.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Hras_surface_colored_by_conservation.png/220px-Hras_surface_colored_by_conservation.png" decoding="async" width="220" height="209" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Hras_surface_colored_by_conservation.png/330px-Hras_surface_colored_by_conservation.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/Hras_surface_colored_by_conservation.png/440px-Hras_surface_colored_by_conservation.png 2x" data-file-width="1300" data-file-height="1236" /></a></span><div class="infobox-caption"><a href="/wiki/HRAS" title="HRAS">HRas</a> structure <a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> 121p, surface colored by conservation in <a href="/wiki/Pfam" title="Pfam">Pfam</a> seed alignment: gold, most conserved; dark cyan, least conserved.</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">Symbol</th><td class="infobox-data" style="background-color: #eee">Ras</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00071">PF00071</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR020849">IPR020849</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/PROSITE" title="PROSITE">PROSITE</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://prosite.expasy.org/PDOC00017">PDOC00017</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Structural_Classification_of_Proteins" class="mw-redirect" title="Structural Classification of Proteins">SCOP2</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=5p21">5p21</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=5p21">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=5p21">SUPFAM</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Conserved_Domain_Database" title="Conserved Domain Database">CDD</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Structure/cdd/cddsrv.cgi?uid=cd04138">cd04138</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">Available protein structures:</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/Pfam" title="Pfam">Pfam</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="http://pfam.xfam.org/family/PF00071?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00071">ECOD</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/Protein_Data_Bank" title="Protein Data Bank">PDB</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.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF00071%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF00071">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF00071">PDBj</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/PDBsum" title="PDBsum">PDBsum</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.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF00071">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p><b>Ras</b>, from "<b>Ra</b>t <b>s</b>arcoma virus", is a <a href="/wiki/Protein_family" title="Protein family">family of related proteins</a> that are expressed in all animal cell lineages and organs. All Ras protein family members belong to a class of protein called <a href="/wiki/Small_GTPase" title="Small GTPase">small GTPase</a>, and are involved in transmitting signals within cells (<a href="/wiki/Cell_(biology)" title="Cell (biology)">cellular</a> <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a>). Ras is the prototypical member of the <a href="/wiki/Ras_superfamily" title="Ras superfamily">Ras superfamily</a> of proteins, which are all related in three-dimensional structure and regulate diverse cell behaviours. </p><p>When Ras is 'switched on' by incoming signals, it subsequently switches on other proteins, which ultimately turn on genes involved in <a href="/wiki/Cell_growth" title="Cell growth">cell growth</a>, <a href="/wiki/Differentiation_(cellular)" class="mw-redirect" title="Differentiation (cellular)">differentiation</a>, and <a href="/wiki/Apoptosis" title="Apoptosis">survival</a>. Mutations in Ras genes can lead to the production of permanently activated Ras proteins, which can cause unintended and overactive signaling inside the cell, even in the absence of incoming signals. </p><p>Because these signals result in cell growth and division, overactive Ras signaling can ultimately lead to <a href="/wiki/Cancer" title="Cancer">cancer</a>.<sup id="cite_ref-pmid10394594_1-0" class="reference"><a href="#cite_note-pmid10394594-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The three Ras genes in humans (<i><a href="/wiki/HRAS" title="HRAS">HRAS</a></i>, <i><a href="/wiki/KRAS" title="KRAS">KRAS</a></i>, and <i><a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a></i>) are the most common <a href="/wiki/Oncogene" title="Oncogene">oncogenes</a> in human cancer; mutations that permanently activate Ras are found in 20 to 25% of all human tumors and up to 90% in certain types of cancer (e.g., <a href="/wiki/Pancreatic_cancer" title="Pancreatic cancer">pancreatic cancer</a>).<sup id="cite_ref-pmid12509763_2-0" class="reference"><a href="#cite_note-pmid12509763-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> For this reason, Ras inhibitors are being studied as a treatment for cancer and other diseases with Ras overexpression. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first two Ras genes, <i><a href="/wiki/HRAS" title="HRAS">HRAS</a></i> and <i><a href="/wiki/KRAS" title="KRAS">KRAS</a></i>, were identified<sup id="cite_ref-pmid12778136_3-0" class="reference"><a href="#cite_note-pmid12778136-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> from studies of two cancer-causing viruses, the Harvey sarcoma virus and Kirsten sarcoma virus, by <a href="/wiki/Edward_M._Scolnick" class="mw-redirect" title="Edward M. Scolnick">Edward M. Scolnick</a> and colleagues at the National Institutes of Health (NIH).<sup id="cite_ref-pmid6289320_4-0" class="reference"><a href="#cite_note-pmid6289320-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> These viruses were discovered originally in rats during the 1960s by Jennifer Harvey<sup id="cite_ref-pmid14243400_5-0" class="reference"><a href="#cite_note-pmid14243400-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Werner_H._Kirsten" title="Werner H. Kirsten">Werner H. Kirsten</a>,<sup id="cite_ref-pmid5538357_6-0" class="reference"><a href="#cite_note-pmid5538357-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> respectively, hence the name <b>Ra</b>t <a href="/wiki/Sarcoma" title="Sarcoma"><b>s</b>arcoma</a>.<sup id="cite_ref-pmid12778136_3-1" class="reference"><a href="#cite_note-pmid12778136-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In 1982, activated and transforming human <i>ras</i> genes were discovered in human cancer cells by <a href="/wiki/Geoffrey_M._Cooper" title="Geoffrey M. Cooper">Geoffrey M. Cooper</a> at Harvard,<sup id="cite_ref-pmid6285471_7-0" class="reference"><a href="#cite_note-pmid6285471-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Mariano_Barbacid" title="Mariano Barbacid">Mariano Barbacid</a> and <a href="/wiki/Stuart_A._Aaronson" title="Stuart A. Aaronson">Stuart A. Aaronson</a> at the NIH,<sup id="cite_ref-pmid6283384_8-0" class="reference"><a href="#cite_note-pmid6283384-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Robert_Weinberg_(biologist)" title="Robert Weinberg (biologist)">Robert Weinberg</a> at MIT,<sup id="cite_ref-pmid6283357_9-0" class="reference"><a href="#cite_note-pmid6283357-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Michael_Wigler" title="Michael Wigler">Michael Wigler</a> at Cold Spring Harbor Laboratory.<sup id="cite_ref-pmid7177195_10-0" class="reference"><a href="#cite_note-pmid7177195-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> A third <i>ras</i> gene was subsequently discovered by researchers in the group of <a href="/wiki/Robin_Weiss" title="Robin Weiss">Robin Weiss</a> at the <a href="/wiki/Institute_of_Cancer_Research" title="Institute of Cancer Research">Institute of Cancer Research</a>,<sup id="cite_ref-pmid6287287_11-0" class="reference"><a href="#cite_note-pmid6287287-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid6304521_12-0" class="reference"><a href="#cite_note-pmid6304521-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Michael_Wigler" title="Michael Wigler">Michael Wigler</a> at Cold Spring Harbor Laboratory,<sup id="cite_ref-pmid6300838_13-0" class="reference"><a href="#cite_note-pmid6300838-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> named <i><a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a></i>, for its initial identification in human neuroblastoma cells. </p><p>The three human <i>ras</i> genes encode extremely similar proteins made up of chains of 188 to 189 amino acids. Their gene symbols are <a href="/wiki/HRAS" title="HRAS">HRAS</a>, <a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a> and <a href="/wiki/KRAS" title="KRAS">KRAS</a>, the latter of which produces the K-Ras4A and K-Ras4B isoforms from alternative <a href="/wiki/Protein_splicing" title="Protein splicing">splicing</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (March 2023)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=2" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Hras_secondary_structure_ribbon.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Hras_secondary_structure_ribbon.png/220px-Hras_secondary_structure_ribbon.png" decoding="async" width="220" height="209" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Hras_secondary_structure_ribbon.png/330px-Hras_secondary_structure_ribbon.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/60/Hras_secondary_structure_ribbon.png/440px-Hras_secondary_structure_ribbon.png 2x" data-file-width="1300" data-file-height="1236" /></a><figcaption><a href="/wiki/HRAS" title="HRAS">HRas</a> structure <a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> 121p, ribbon showing strands in purple, helices in aqua, loops in gray. Also shown are the bound GTP analog and magnesium ion.</figcaption></figure> <p>Ras contains six <a href="/wiki/Beta_strand" class="mw-redirect" title="Beta strand">beta strands</a> and five <a href="/wiki/Alpha_helices" class="mw-redirect" title="Alpha helices">alpha helices</a>.<sup id="cite_ref-pmid11701921_14-0" class="reference"><a href="#cite_note-pmid11701921-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> It consists of two domains: a G domain of 166 <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> (about 20 kDa) that binds guanosine nucleotides, and a C-terminal membrane targeting region (CAAX-COOH, also known as <a href="/w/index.php?title=CAAX_box&action=edit&redlink=1" class="new" title="CAAX box (page does not exist)">CAAX box</a>), which is lipid-modified by <a href="/wiki/Farnesyl_transferase" class="mw-redirect" title="Farnesyl transferase">farnesyl transferase</a>, <a href="/wiki/RCE1" title="RCE1">RCE1</a>, and <a href="/wiki/ICMT" title="ICMT">ICMT</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (March 2023)">citation needed</span></a></i>]</sup> </p><p>The G domain contains five G motifs that bind GDP/GTP directly. The G1 motif, or the P-loop, binds the beta phosphate of GDP and GTP. The G2 motif, also called Switch I or SW1, contains threonine35, which binds the terminal phosphate (γ-phosphate) of GTP and the divalent magnesium ion bound in the active site. The G3 motif, also called Switch II or SW2, has a DXXGQ motif. The D is aspartate57, which is specific for guanine versus adenine binding, and Q is glutamine61, the crucial residue that activates a catalytic water molecule for hydrolysis of GTP to GDP. The G4 motif contains a LVGNKxDL motif, and provides specific interaction to guanine. The G5 motif contains a SAK consensus sequence. The A is alanine146, which provides specificity for guanine rather than adenine. </p><p>The two switch motifs, G2 (SW1) and G3 (SW2), are the main parts of the protein that move when GTP is hydrolyzed into GDP. This conformational change by the two switch motifs is what mediates the basic functionality as a molecular switch protein. This GTP-bound state of Ras is the "on" state, and the GDP-bound state is the "off" state. The two switch motifs have a number of conformations when binding GTP or GDP or no nucleotide (when bound to SOS1, which releases the nucleotide).<sup id="cite_ref-RasCore_15-0" class="reference"><a href="#cite_note-RasCore-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Ras also binds a <a href="/wiki/Magnesium_in_biology" title="Magnesium in biology">magnesium ion</a> which helps to coordinate nucleotide binding. </p> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=3" title="Edit section: Function"><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:Signal_transduction_pathways.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/300px-Signal_transduction_pathways.svg.png" decoding="async" width="300" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/450px-Signal_transduction_pathways.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/600px-Signal_transduction_pathways.svg.png 2x" data-file-width="1858" data-file-height="1364" /></a><figcaption>Overview of signal transduction pathways involved in <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a></figcaption></figure> <p>Ras proteins function as binary molecular switches that control intracellular signaling networks. Ras-regulated <a href="/wiki/Signal_transduction" title="Signal transduction">signal pathways</a> control such processes as actin <a href="/wiki/Cytoskeletal" class="mw-redirect" title="Cytoskeletal">cytoskeletal</a> integrity, <a href="/wiki/Cell_proliferation" title="Cell proliferation">cell proliferation</a>, <a href="/wiki/Cell_differentiation" class="mw-redirect" title="Cell differentiation">cell differentiation</a>, <a href="/wiki/Cell_adhesion" title="Cell adhesion">cell adhesion</a>, <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, and <a href="/wiki/Cell_migration" title="Cell migration">cell migration</a>. Ras and Ras-related proteins are often deregulated in cancers, leading to increased invasion and <a href="/wiki/Metastasis" title="Metastasis">metastasis</a>, and decreased apoptosis. </p><p>Ras activates several pathways, of which the <a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">mitogen-activated protein (MAP) kinase cascade</a> has been well-studied. This cascade transmits signals downstream and results in the <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of genes involved in cell growth and division.<sup id="cite_ref-Lodish_16-0" class="reference"><a href="#cite_note-Lodish-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Another Ras-activated signaling pathway is the <a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a>, which stimulates protein synthesis, cellular migration and growth, and inhibits apoptosis.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Activation_and_deactivation">Activation and deactivation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=4" title="Edit section: Activation and deactivation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ras is a <a href="/wiki/Guanosine" title="Guanosine">guanosine</a>-<a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a>-binding protein. Specifically, it is a single-subunit <a href="/wiki/Small_GTPase" title="Small GTPase">small GTPase</a>, which is related in structure to the G<sub>α</sub> subunit of <a href="/wiki/Heterotrimeric_G_proteins" class="mw-redirect" title="Heterotrimeric G proteins">heterotrimeric G proteins</a> (large GTPases). G proteins function as binary signaling switches with "on" and "off" states. In the "off" state it is bound to the nucleotide <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">guanosine diphosphate</a> (GDP), while in the "on" state, Ras is bound to <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">guanosine triphosphate</a> (GTP), which has an extra <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> group as compared to GDP. This extra phosphate holds the two switch regions in a "loaded-spring" configuration (specifically the Thr-35 and Gly-60). When released, the switch regions relax which causes a <a href="/wiki/Conformational_change" title="Conformational change">conformational change</a> into the inactive state. Hence, activation and deactivation of Ras and other small G proteins are controlled by cycling between the active GTP-bound and inactive GDP-bound forms. </p><p>The process of exchanging the bound nucleotide is facilitated by <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">guanine nucleotide exchange factors</a> (GEFs) and <a href="/wiki/GTPase_activating_proteins" class="mw-redirect" title="GTPase activating proteins">GTPase activating proteins</a> (GAPs). As per its classification, Ras has an intrinsic <a href="/wiki/GTPase" title="GTPase">GTPase</a> activity, which means that the protein on its own will <a href="/wiki/Hydrolyze" class="mw-redirect" title="Hydrolyze">hydrolyze</a> a bound GTP molecule into GDP. However this process is too slow for efficient function, and hence the GAP for Ras, RasGAP, may bind to and stabilize the catalytic machinery of Ras, supplying additional catalytic residues ("<a href="/wiki/Arginine_finger" title="Arginine finger">arginine finger</a>") such that a water molecule is optimally positioned for <a href="/wiki/Nucleophilic" class="mw-redirect" title="Nucleophilic">nucleophilic</a> attack on the gamma-phosphate of GTP. An inorganic phosphate is released and the Ras molecule is now bound to a GDP. Since the GDP-bound form is "off" or "inactive" for signaling, GTPase Activating Protein <i>inactivates</i> Ras by activating its GTPase activity. Thus, GAPs accelerate Ras <i>inactivation</i>. </p><p>GEFs catalyze a "push and pull" reaction which releases GDP from Ras. They insert close to the P-loop and magnesium <a href="/wiki/Cation" class="mw-redirect" title="Cation">cation</a> binding site and inhibit the interaction of these with the gamma phosphate <a href="/wiki/Anion" class="mw-redirect" title="Anion">anion</a>. Acidic (negative) residues in switch II "pull" a lysine in the P-loop away from the GDP which "pushes" switch I away from the guanine. The contacts holding GDP in place are broken and it is released into the cytoplasm. Because intracellular GTP is abundant relative to GDP (approximately 10 fold more)<sup id="cite_ref-Lodish_16-1" class="reference"><a href="#cite_note-Lodish-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> GTP predominantly re-enters the nucleotide binding pocket of Ras and reloads the spring. Thus GEFs facilitate Ras <i>activation</i>.<sup id="cite_ref-pmid11701921_14-1" class="reference"><a href="#cite_note-pmid11701921-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Well known GEFs include <a href="/wiki/Son_of_Sevenless" title="Son of Sevenless">Son of Sevenless</a> (Sos) and <a href="/wiki/Cdc25" title="Cdc25">cdc25</a> which include the <a href="/wiki/RasGEF_domain" title="RasGEF domain">RasGEF domain</a>. </p><p>The balance between GEF and GAP activity determines the guanine nucleotide status of Ras, thereby regulating Ras activity. </p><p>In the GTP-bound conformation, Ras has a high affinity for numerous <a href="/wiki/Effector_(biology)" title="Effector (biology)">effectors</a> which allow it to carry out its functions. These include <a href="/wiki/PI3K" class="mw-redirect" title="PI3K">PI3K</a>. Other small GTPases may bind adaptors such as <a href="/wiki/Arfaptin" title="Arfaptin">arfaptin</a> or second messenger systems such as <a href="/wiki/Adenylyl_cyclase" title="Adenylyl cyclase">adenylyl cyclase</a>. The Ras binding domain is found in many effectors and invariably binds to one of the switch regions, because these change conformation between the active and inactive forms. However, they may also bind to the rest of the protein surface. </p><p>Other proteins exist that may change the activity of Ras family proteins. One example is GDI (GDP Disassociation Inhibitor). These function by slowing the exchange of GDP for GTP, thus prolonging the inactive state of Ras family members. Other proteins that augment this cycle may exist. </p> <div class="mw-heading mw-heading3"><h3 id="Membrane_attachment">Membrane attachment</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=5" title="Edit section: Membrane attachment"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ras is attached to the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> owing to its <a href="/wiki/Prenylation" title="Prenylation">prenylation</a> and <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> (<a href="/wiki/HRAS" title="HRAS">HRAS</a> and <a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a>) or the combination of <a href="/wiki/Prenylation" title="Prenylation">prenylation</a> and a polybasic sequence adjacent to the prenylation site (<a href="/wiki/KRAS" title="KRAS">KRAS</a>). The C-terminal <a href="/w/index.php?title=CaaX_box&action=edit&redlink=1" class="new" title="CaaX box (page does not exist)">CaaX box</a> of Ras first gets farnesylated at its Cys residue in the cytosol, allowing Ras to loosely insert into the membrane of the <a href="/wiki/Endoplasmatic_reticulum" class="mw-redirect" title="Endoplasmatic reticulum">endoplasmatic reticulum</a> and other cellular membranes. The Tripeptide (aaX) is then cleaved from the C-terminus by a specific prenyl-protein specific endoprotease and the new C-terminus is methylated by a <a href="/wiki/Methyltransferase" title="Methyltransferase">methyltransferase</a>. KRas processing is completed at this stage. Dynamic electrostatic interactions between its positively charged basic sequence with negative charges at the inner leaflet of the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> account for its predominant localization at the cell surface at steady-state. <a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a> and <a href="/wiki/HRAS" title="HRAS">HRAS</a> are further processed on the surface of the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> by <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> of one or two Cys residues, respectively, adjacent to the <a href="/w/index.php?title=CaaX_box&action=edit&redlink=1" class="new" title="CaaX box (page does not exist)">CaaX box</a>. The proteins thereby become stably membrane anchored (lipid-rafts) and are transported to the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> on <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a> of the <a href="/wiki/Secretory_pathway" class="mw-redirect" title="Secretory pathway">secretory pathway</a>. Depalmitoylation by <a href="/wiki/Acyl-protein_thioesterase" title="Acyl-protein thioesterase">acyl-protein thioesterases</a> eventually releases the proteins from the membrane, allowing them to enter another cycle of palmitoylation and depalmitoylation.<sup id="cite_ref-Rocks_19-0" class="reference"><a href="#cite_note-Rocks-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> This cycle is believed to prevent the leakage of <a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a> and <a href="/wiki/HRAS" title="HRAS">HRAS</a> to other membranes over time and to maintain their steady-state localization along the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a>, <a href="/wiki/Secretory_pathway" class="mw-redirect" title="Secretory pathway">secretory pathway</a>, <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> and inter-linked <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a> pathway. </p> <div class="mw-heading mw-heading2"><h2 id="Members">Members</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=6" title="Edit section: Members"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The clinically most notable members of the Ras subfamily are <a href="/wiki/HRAS" title="HRAS">HRAS</a>, <a href="/wiki/KRAS" title="KRAS">KRAS</a> and <a href="/wiki/Neuroblastoma_RAS_viral_oncogene_homolog" title="Neuroblastoma RAS viral oncogene homolog">NRAS</a>, mainly for being implicated in many types of cancer.<sup id="cite_ref-bos_20-0" class="reference"><a href="#cite_note-bos-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>However, there are many other members of this subfamily as well:<sup id="cite_ref-pmid15731001_21-0" class="reference"><a href="#cite_note-pmid15731001-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> <a href="/wiki/DIRAS1" title="DIRAS1">DIRAS1</a>; <a href="/w/index.php?title=DIRAS2&action=edit&redlink=1" class="new" title="DIRAS2 (page does not exist)">DIRAS2</a>; <a href="/wiki/DIRAS3" class="mw-redirect" title="DIRAS3">DIRAS3</a>; <a href="/w/index.php?title=E-Ras&action=edit&redlink=1" class="new" title="E-Ras (page does not exist)">ERAS</a>; <a href="/wiki/GEM_(gene)" title="GEM (gene)">GEM</a>; <a href="/wiki/MRAS" title="MRAS">MRAS</a>; <a href="/w/index.php?title=NKIRAS1&action=edit&redlink=1" class="new" title="NKIRAS1 (page does not exist)">NKIRAS1</a>; <a href="/wiki/NKIRAS2" title="NKIRAS2">NKIRAS2</a>; <a href="/wiki/RALA" title="RALA">RALA</a>; <a href="/wiki/RALB" title="RALB">RALB</a>; <a href="/wiki/RAP1A" title="RAP1A">RAP1A</a>; <a href="/wiki/RAP1B" title="RAP1B">RAP1B</a>; <a href="/wiki/RAP2A" title="RAP2A">RAP2A</a>; <a href="/wiki/RAP2B" title="RAP2B">RAP2B</a>; <a href="/w/index.php?title=RAP2C&action=edit&redlink=1" class="new" title="RAP2C (page does not exist)">RAP2C</a>; <a href="/wiki/RASD1" title="RASD1">RASD1</a>; <a href="/wiki/RASD2" title="RASD2">RASD2</a>; <a href="/w/index.php?title=RASL10A&action=edit&redlink=1" class="new" title="RASL10A (page does not exist)">RASL10A</a>; <a href="/w/index.php?title=RASL10B&action=edit&redlink=1" class="new" title="RASL10B (page does not exist)">RASL10B</a>; <a href="/w/index.php?title=RASL11A&action=edit&redlink=1" class="new" title="RASL11A (page does not exist)">RASL11A</a>; <a href="/wiki/RASL11B" title="RASL11B">RASL11B</a>; <a href="/w/index.php?title=RASL12&action=edit&redlink=1" class="new" title="RASL12 (page does not exist)">RASL12</a>; <a href="/w/index.php?title=REM1&action=edit&redlink=1" class="new" title="REM1 (page does not exist)">REM1</a>; <a href="/w/index.php?title=REM2&action=edit&redlink=1" class="new" title="REM2 (page does not exist)">REM2</a>; <a href="/wiki/RERG" title="RERG">RERG</a>; <a href="/w/index.php?title=RERGL&action=edit&redlink=1" class="new" title="RERGL (page does not exist)">RERGL</a>; <a href="/wiki/RRAD_(gene)" title="RRAD (gene)">RRAD</a>; <a href="/wiki/RRAS" title="RRAS">RRAS</a>; <a href="/wiki/RRAS2" title="RRAS2">RRAS2</a> </p> <div class="mw-heading mw-heading2"><h2 id="Ras_in_cancer">Ras in cancer</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=7" title="Edit section: Ras in cancer"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mutations in the Ras family of <a href="/wiki/Proto-oncogene" class="mw-redirect" title="Proto-oncogene">proto-oncogenes</a> (comprising H-Ras, N-Ras and K-Ras) are very common, being found in 20% to 30% of all human tumors.<sup id="cite_ref-bos_20-1" class="reference"><a href="#cite_note-bos-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> It is reasonable to speculate that a pharmacological approach that curtails Ras activity may represent a possible method to inhibit certain cancer types. Ras point mutations are the single most common abnormality of human proto-oncogenes.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> <a href="/w/index.php?title=Ras_inhibitor&action=edit&redlink=1" class="new" title="Ras inhibitor (page does not exist)">Ras inhibitor</a> trans-farnesylthiosalicylic acid (FTS, <a href="/w/index.php?title=Salirasib&action=edit&redlink=1" class="new" title="Salirasib (page does not exist)">Salirasib</a>) exhibits profound anti-oncogenic effects in many cancer cell lines.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inappropriate_activation">Inappropriate activation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=8" title="Edit section: Inappropriate activation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Inappropriate activation of the gene has been shown to play a key role in improper signal transduction, proliferation and malignant transformation.<sup id="cite_ref-Lodish_16-2" class="reference"><a href="#cite_note-Lodish-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Mutations in a number of different genes as well as RAS itself can have this effect. <a href="/wiki/Oncogene" title="Oncogene">Oncogenes</a> such as p210BCR-ABL or the growth receptor erbB are upstream of Ras, so if they are constitutively activated their signals will transduce through Ras.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (March 2023)">citation needed</span></a></i>]</sup> </p><p>The <a href="/wiki/Tumour_suppressor_gene" class="mw-redirect" title="Tumour suppressor gene">tumour suppressor gene</a> <a href="/wiki/Neurofibromin_1" class="mw-redirect" title="Neurofibromin 1">NF1</a> encodes a Ras-GAP – its mutation in <a href="/wiki/Neurofibromatosis" title="Neurofibromatosis">neurofibromatosis</a> will mean that Ras is less likely to be inactivated. Ras can also be amplified, although this only occurs occasionally in tumours. </p><p>Finally, Ras oncogenes can be activated by point mutations so that the GTPase reaction can no longer be stimulated by GAP – this increases the half life of active Ras-GTP mutants.<sup id="cite_ref-Reuter_25-0" class="reference"><a href="#cite_note-Reuter-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Constitutively_active_Ras">Constitutively active Ras</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=9" title="Edit section: Constitutively active Ras"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Constitutively active Ras</i> (<i>Ras<sup>D</sup></i>) is one which contains mutations that prevent GTP hydrolysis, thus locking Ras in a permanently 'On' state. </p><p>The most common mutations are found at residue G12 in the <a href="/wiki/P-loop" class="mw-redirect" title="P-loop">P-loop</a> and the catalytic residue Q61. </p> <ul><li>The glycine to valine mutation at <i>residue 12</i> renders the GTPase domain of Ras insensitive to inactivation by GAP and thus stuck in the "on state". Ras requires a GAP for inactivation as it is a relatively poor catalyst on its own, as opposed to other G-domain-containing proteins such as the alpha subunit of heterotrimeric G proteins.</li> <li><i>Residue 61</i><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> is responsible for stabilizing the transition state for GTP hydrolysis. Because enzyme catalysis in general is achieved by lowering the energy barrier between substrate and product, mutation of Q61 to K (Glutamine to Lysine) necessarily reduces the rate of intrinsic Ras GTP hydrolysis to physiologically meaningless levels.</li></ul> <p>See also "dominant negative" mutants such as S17N and D119N. </p> <div class="mw-heading mw-heading3"><h3 id="Ras-targeted_cancer_treatments">Ras-targeted cancer treatments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=10" title="Edit section: Ras-targeted cancer treatments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Reovirus" class="mw-redirect" title="Reovirus">Reovirus</a> was noted to be a potential cancer therapeutic when studies suggested it reproduces well in certain cancer cell lines. It replicates specifically in cells that have an activated Ras pathway (a cellular signaling pathway that is involved in cell growth and differentiation).<sup id="cite_ref-pmid19806501_27-0" class="reference"><a href="#cite_note-pmid19806501-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Reovirus replicates in and eventually kills Ras-activated tumour cells and as cell death occurs, progeny virus particles are free to infect surrounding cancer cells. This cycle of infection, replication and cell death is believed to be repeated until all tumour cells carrying an activated Ras pathway are destroyed.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (March 2023)">citation needed</span></a></i>]</sup> </p><p>Another tumor-lysing virus that specifically targets tumor cells with an activated Ras pathway is a type II <a href="/wiki/Herpes_simplex_virus" title="Herpes simplex virus">herpes simplex virus</a> (HSV-2) based agent, designated FusOn-H2.<sup id="cite_ref-pmid16569513_28-0" class="reference"><a href="#cite_note-pmid16569513-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Activating mutations of the Ras protein and upstream elements of the Ras protein may play a role in more than two-thirds of all human cancers, including most metastatic disease. <a href="/wiki/Reolysin" class="mw-redirect" title="Reolysin">Reolysin</a>, a formulation of reovirus, and FusOn-H2 are currently in clinical trials or under development for the treatment of various cancers.<sup id="cite_ref-pmid19565924_29-0" class="reference"><a href="#cite_note-pmid19565924-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In addition, a treatment based on siRNA anti-mutated <a href="/wiki/KRAS" title="KRAS">K-RAS</a> (G12D) called siG12D LODER is currently in clinical trials for the treatment of locally advanced pancreatic cancer (NCT01188785, NCT01676259).<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><p>In <a href="/wiki/Glioblastoma" title="Glioblastoma">glioblastoma</a> mouse models SHP2 levels were heightened in cancerous brain cells. Inhibiting <a href="/wiki/PTPN11" title="PTPN11">SHP2</a> in turn inhibited Ras dephosphorylation. This reduced tumor sizes and accompanying rise in survival rates.<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><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> </p><p>Other strategies have attempted to manipulate the regulation of the above-mentioned localization of Ras. <a href="/wiki/Farnesyltransferase_inhibitor" title="Farnesyltransferase inhibitor">Farnesyltransferase inhibitors</a> have been developed to stop the <a href="/wiki/Farnesylation" class="mw-redirect" title="Farnesylation">farnesylation</a> of Ras and therefore weaken its affinity to membranes.<sup id="cite_ref-pmid12509763_2-1" class="reference"><a href="#cite_note-pmid12509763-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Other inhibitors are targeting the <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> cycle of Ras through inhibiting depalmitoylation by <a href="/wiki/Acyl-protein_thioesterase#Inhibition_of_acyl-protein_thioesterases" title="Acyl-protein thioesterase">acyl-protein thioesterases</a>, potentially leading to a destabilization of the Ras cycle.<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> </p><p>A novel inhibitor finding strategy for mutated Ras molecules was described in.<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> The Ras mutations in the 12th residue position inhibit the bound of the regulatory GAP molecule to the mutated Ras, causing uncontrolled cell growth. The novel strategy proposes finding small glue molecules, which attach the mutated Ras to the GAP, prohibiting uncontrolled cell growth and restoring the normal function. For this goal a theoretical Ras-GAP conformation was designed with a several Å gap between the molecules, and a high-throughput in silico docking was performed for finding gluing agents. As a proof of concept, two novel molecules were described with satisfying biological activity. </p> <div class="mw-heading mw-heading2"><h2 id="In_other_species">In other species</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=11" title="Edit section: In other species"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In most of the cell types of most species, most <i>Ras</i> is the GDP type. This is true for <i><a href="/wiki/Xenopus" title="Xenopus">Xenopus</a></i> <a href="/wiki/Oocyte" title="Oocyte">oocytes</a> and <a href="/wiki/Mouse" title="Mouse">mouse</a> <a href="/wiki/Fibroblast" title="Fibroblast">fibroblasts</a>.<sup id="cite_ref-Bollag-McCormick-1991_35-0" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Xenopus_laevis"><i>Xenopus laevis</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=12" title="Edit section: Xenopus laevis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As mentioned above most <i>X.</i> oocyte <i>Ras</i> is the GDP conjugate. Mammal <i>Ras</i> induces <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> in <i><a href="/wiki/Xenopus_laevis" class="mw-redirect" title="Xenopus laevis">X. laevis</a></i> oocytes almost certainly by potentiating <a href="/wiki/Insulin" title="Insulin">insulin</a>-induced meiosis, but not <a href="/wiki/Progesterone" title="Progesterone">progesterone</a>-induced. <a href="/wiki/Protein_synthesis" class="mw-redirect" title="Protein synthesis">Protein synthesis</a> does not seem to be a part of this step. Injection increases synthesis of <a href="/wiki/Diacylglycerol" class="mw-redirect" title="Diacylglycerol">diacylglycerol</a> from <a href="/wiki/Phosphatidylcholine" title="Phosphatidylcholine">phosphatidylcholine</a>. Some meiosis effects are <a href="/wiki/Physiological_antagonist" class="mw-redirect" title="Physiological antagonist">antagonized</a> by <i><a href="/wiki/Rap1" title="Rap1">rap1</a></i> (and by a <i>Ras</i> modified to dock incorrectly). Both <i>rap1</i> and the modified <i>Ras</i> are co-antagonists with <a href="/wiki/P120Ras_GAP" class="mw-redirect" title="P120Ras GAP">p120Ras GAP</a> in this pathway.<sup id="cite_ref-Bollag-McCormick-1991_35-1" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Drosophila_melanogaster"><i>Drosophila melanogaster</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=13" title="Edit section: Drosophila melanogaster"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Expressed in all tissues of <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> but mostly in neural cells. Overexpression is somewhat lethal and, during development, produces eye and wing abnormalities. (This parallels - and may be the reason for - similar abnormalities due to mutated <a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">receptor tyrosine kinases</a>.) The <i>D.</i> genes for <i>ras</i>es in mammals produce abnormalities.<sup id="cite_ref-Bollag-McCormick-1991_35-2" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Aplysia"><i>Aplysia</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=14" title="Edit section: Aplysia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most expression in <i><a href="/wiki/Aplysia" title="Aplysia">Aplysia</a></i> spp. is in neural cells.<sup id="cite_ref-Bollag-McCormick-1991_35-3" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Caenorhabditis_elegans"><i>Caenorhabditis elegans</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=15" title="Edit section: Caenorhabditis elegans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><span class="anchor" id="let_60"></span>The gene in <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i> is <i>let 60</i>. Also appears to play a role in receptor tyrosine kinase formation in this model. Overexpression yields a multivulval development due to its involvement in that region's normal development; overexpression in effector sites in lethal.<sup id="cite_ref-Bollag-McCormick-1991_35-4" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dictyostelium_discoideum"><i>Dictyostelium discoideum</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=16" title="Edit section: Dictyostelium discoideum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Essential in <i><a href="/wiki/Dictyostelium_discoideum" title="Dictyostelium discoideum">Dictyostelium discoideum</a></i>. This is evidenced by severe developmental failure in deficient <i>ras</i> expression and by significant impairment of various life activities when artificially expressed, such as: increased concentration of <a href="/wiki/Inositol_phosphate" title="Inositol phosphate">inositol phosphates</a>; likely reduction of <a href="/wiki/Cyclic_adenosine_monophosphate" title="Cyclic adenosine monophosphate">cAMP</a> binding to <a href="/wiki/Chemotaxis" title="Chemotaxis">chemotaxis</a> receptors; and that is likely the reason <a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cGMP</a> synthesis is impaired. <a href="/wiki/Adenylate_cyclase" class="mw-redirect" title="Adenylate cyclase">Adenylate cyclase</a> activity is unaffected by <i>ras</i>.<sup id="cite_ref-Bollag-McCormick-1991_35-5" class="reference"><a href="#cite_note-Bollag-McCormick-1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <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=Ras_GTPase&action=edit&section=17" 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 reflist-columns-2"> <ol class="references"> <li id="cite_note-pmid10394594-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid10394594_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 (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFGoodsell1999" class="citation journal cs1">Goodsell DS (1999). <a rel="nofollow" class="external text" href="http://theoncologist.alphamedpress.org/cgi/content/full/4/3/263">"The molecular perspective: the ras oncogene"</a>. <i>The Oncologist</i>. <b>4</b> (3): 263–4. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1634%2Ftheoncologist.4-3-263">10.1634/theoncologist.4-3-263</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/10394594">10394594</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Oncologist&rft.atitle=The+molecular+perspective%3A+the+ras+oncogene&rft.volume=4&rft.issue=3&rft.pages=263-4&rft.date=1999&rft_id=info%3Adoi%2F10.1634%2Ftheoncologist.4-3-263&rft_id=info%3Apmid%2F10394594&rft.aulast=Goodsell&rft.aufirst=DS&rft_id=http%3A%2F%2Ftheoncologist.alphamedpress.org%2Fcgi%2Fcontent%2Ffull%2F4%2F3%2F263&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" class="Z3988"></span></span> </li> <li id="cite_note-pmid12509763-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid12509763_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid12509763_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDownward2003" class="citation journal cs1">Downward J (January 2003). 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Cancer</i>. <b>3</b> (1): 11–22. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrc969">10.1038/nrc969</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/12509763">12509763</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:43074411">43074411</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews.+Cancer&rft.atitle=Targeting+RAS+signalling+pathways+in+cancer+therapy&rft.volume=3&rft.issue=1&rft.pages=11-22&rft.date=2003-01&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A43074411%23id-name%3DS2CID&rft_id=info%3Apmid%2F12509763&rft_id=info%3Adoi%2F10.1038%2Fnrc969&rft.aulast=Downward&rft.aufirst=J&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" class="Z3988"></span></span> </li> <li id="cite_note-pmid12778136-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid12778136_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid12778136_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMalumbresBarbacid2003" class="citation journal cs1">Malumbres M, Barbacid M (June 2003). 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Cancer</i>. <b>3</b> (6): 459–65. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrc1097">10.1038/nrc1097</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/12778136">12778136</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:27928171">27928171</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews.+Cancer&rft.atitle=RAS+oncogenes%3A+the+first+30+years&rft.volume=3&rft.issue=6&rft.pages=459-65&rft.date=2003-06&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A27928171%23id-name%3DS2CID&rft_id=info%3Apmid%2F12778136&rft_id=info%3Adoi%2F10.1038%2Fnrc1097&rft.aulast=Malumbres&rft.aufirst=M&rft.au=Barbacid%2C+M&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" class="Z3988"></span></span> </li> <li id="cite_note-pmid6289320-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid6289320_4-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChangGondaEllisScolnick1982" class="citation journal cs1">Chang EH, Gonda MA, Ellis RW, Scolnick EM, Lowy DR (August 1982). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC346782">"Human genome contains four genes homologous to transforming genes of Harvey and Kirsten murine sarcoma viruses"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>79</b> (16): 4848–52. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1982PNAS...79.4848C">1982PNAS...79.4848C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.79.16.4848">10.1073/pnas.79.16.4848</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC346782">346782</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/6289320">6289320</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Human+genome+contains+four+genes+homologous+to+transforming+genes+of+Harvey+and+Kirsten+murine+sarcoma+viruses&rft.volume=79&rft.issue=16&rft.pages=4848-52&rft.date=1982-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC346782%23id-name%3DPMC&rft_id=info%3Apmid%2F6289320&rft_id=info%3Adoi%2F10.1073%2Fpnas.79.16.4848&rft_id=info%3Abibcode%2F1982PNAS...79.4848C&rft.aulast=Chang&rft.aufirst=EH&rft.au=Gonda%2C+MA&rft.au=Ellis%2C+RW&rft.au=Scolnick%2C+EM&rft.au=Lowy%2C+DR&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC346782&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" class="Z3988"></span></span> </li> <li id="cite_note-pmid14243400-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid14243400_5-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarvey1964" class="citation journal cs1">Harvey JJ (December 1964). 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"Regulators and effectors of ras proteins". <i>Annual Review of Cell Biology</i>. <b>7</b> (1). <a href="/wiki/Annual_Reviews_(publisher)" title="Annual Reviews (publisher)">Annual Reviews</a>: 601–32. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.cb.07.110191.003125">10.1146/annurev.cb.07.110191.003125</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/1667084">1667084</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Cell+Biology&rft.atitle=Regulators+and+effectors+of+ras+proteins&rft.volume=7&rft.issue=1&rft.pages=601-32&rft.date=1991&rft_id=info%3Adoi%2F10.1146%2Fannurev.cb.07.110191.003125&rft_id=info%3Apmid%2F1667084&rft.aulast=Bollag&rft.aufirst=G&rft.au=McCormick%2C+F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" 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=Ras_GTPase&action=edit&section=18" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAgrawalSomani2009" class="citation journal cs1">Agrawal AG, Somani RR (June 2009). "Farnesyltransferase inhibitor as anticancer agent". <i>Mini Reviews in Medicinal Chemistry</i>. <b>9</b> (6): 638–52. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2174%2F138955709788452702">10.2174/138955709788452702</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/19519490">19519490</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Mini+Reviews+in+Medicinal+Chemistry&rft.atitle=Farnesyltransferase+inhibitor+as+anticancer+agent&rft.volume=9&rft.issue=6&rft.pages=638-52&rft.date=2009-06&rft_id=info%3Adoi%2F10.2174%2F138955709788452702&rft_id=info%3Apmid%2F19519490&rft.aulast=Agrawal&rft.aufirst=AG&rft.au=Somani%2C+RR&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARas+GTPase" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ras_GTPase&action=edit&section=19" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080927031424/http://www.ncic.cancer.ca/ncic/internet/mediareleaselist/0,,84658243_85783616_88058948_langId-en.html">"Brain tumour findings offer hope of new strategy Canadian Cancer Society says" at ncic.cancer.ca</a></li> <li><a rel="nofollow" class="external text" href="https://arstechnica.com/uncategorized/2003/11/3072-2/">"Novel cancer treatment gets NCI support" at arstechnica.com</a></li> <li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=ras+Proteins">ras+Proteins</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=ras+Genes">ras+Genes</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><a rel="nofollow" class="external text" href="http://www.sdbonline.org/fly/torstoll/ras85-1.htm"><i>Drosophila</i> <i>Ras oncogene at 85D</i> - The Interactive Fly</a></li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=iWzfp9d0pw4">"Animation of ras activation by EGFR"</a></li> <li><a rel="nofollow" class="external text" href="http://dunbrack.fccc.edu/rascore">"Rascore: A tool for analyzing RAS protein structures"</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist 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abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Intracellular_signaling_peptides_and_proteins" title="Template:Intracellular signaling peptides and proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Intracellular_signaling_peptides_and_proteins" title="Template talk:Intracellular signaling peptides and proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Intracellular_signaling_peptides_and_proteins" title="Special:EditPage/Template:Intracellular signaling peptides and proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Intracellular_signaling_peptides_and_proteins" style="font-size:114%;margin:0 4em"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular signaling peptides and proteins</a></div></th></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">MAP</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><i>see <a href="/wiki/Template:MAP_kinase_pathway" title="Template:MAP kinase pathway">MAP kinase pathway</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Calcium</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/Intracellular_calcium-sensing_proteins" title="Intracellular calcium-sensing proteins">Intracellular calcium-sensing proteins</a></li> <li><a href="/wiki/Calcineurin" title="Calcineurin">Calcineurin</a></li> <li><a href="/wiki/Ca2%2B/calmodulin-dependent_protein_kinase" class="mw-redirect" title="Ca2+/calmodulin-dependent protein kinase">Ca2+/calmodulin-dependent protein kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%"><a href="/wiki/G_protein" title="G protein">G protein</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/Heterotrimeric_G_protein" title="Heterotrimeric G protein">Heterotrimeric</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP</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/Heterotrimeric_G_protein" title="Heterotrimeric G protein">Heterotrimeric G protein</a> <ul><li><a href="/wiki/Gs_alpha_subunit" title="Gs alpha subunit">Gs</a>/<a href="/wiki/Gi_alpha_subunit" title="Gi alpha subunit">Gi</a></li></ul></li> <li><a href="/wiki/Adenylate_cyclase" class="mw-redirect" title="Adenylate cyclase">Adenylate cyclase</a></li> <li><a href="/wiki/Cyclic_adenosine_monophosphate" title="Cyclic adenosine monophosphate">cAMP</a></li> <li><a href="/wiki/3%27,5%27-cyclic-AMP_phosphodiesterase" title="3',5'-cyclic-AMP phosphodiesterase">3',5'-cyclic-AMP phosphodiesterase</a></li> <li><a href="/wiki/Protein_kinase_A" title="Protein kinase A">Protein kinase A</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cGMP</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/Transducin" title="Transducin">Transducin</a></li> <li><a href="/wiki/Gustducin" title="Gustducin">Gustducin</a></li> <li><a href="/wiki/Guanylate_cyclase" title="Guanylate cyclase">Guanylate cyclase</a></li> <li><a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cGMP</a></li> <li><a href="/wiki/3%27,5%27-cyclic-GMP_phosphodiesterase" title="3',5'-cyclic-GMP phosphodiesterase">3',5'-cyclic-GMP phosphodiesterase</a></li> <li><a href="/wiki/CGMP-dependent_protein_kinase" title="CGMP-dependent protein kinase">Protein kinase G</a></li></ul> </div></td></tr><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/G_alpha_subunit" title="G alpha subunit">G alpha subunit</a> G<sub>α</sub> <ul><li><a href="/wiki/GNAO1" title="GNAO1">GNAO1</a></li> <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/GNAT1" title="GNAT1">GNAT1</a></li> <li><a href="/wiki/GNAT2" title="GNAT2">GNAT2</a></li> <li><a href="/wiki/GNAT3" title="GNAT3">GNAT3</a></li> <li><a href="/wiki/GNAZ" title="GNAZ">GNAZ</a></li> <li><a href="/wiki/GNAS_complex_locus" title="GNAS complex locus">GNAS</a></li> <li><a href="/wiki/GNAL" title="GNAL">GNAL</a></li> <li><a href="/wiki/GNAQ" title="GNAQ">GNAQ</a></li> <li><a href="/wiki/GNA11" title="GNA11">GNA11</a></li> <li><a href="/wiki/GNA12" title="GNA12">GNA12</a></li> <li><a href="/wiki/GNA13" title="GNA13">GNA13</a></li> <li><a href="/wiki/GNA14" title="GNA14">GNA14</a></li> <li><a href="/w/index.php?title=GNA15/GNA16&action=edit&redlink=1" class="new" title="GNA15/GNA16 (page does not exist)">GNA15/GNA16</a></li></ul></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/G_beta-gamma_complex" title="G beta-gamma complex">G beta-gamma complex</a> G<sub>β</sub> <ul><li><a href="/wiki/GNB1" title="GNB1">GNB1</a></li> <li><a href="/wiki/GNB2" title="GNB2">GNB2</a></li> <li><a href="/wiki/GNB3" title="GNB3">GNB3</a></li> <li><a href="/wiki/GNB4" title="GNB4">GNB4</a></li> <li><a href="/wiki/GNB5" title="GNB5">GNB5</a></li></ul></li> <li>G<sub>γ</sub> <ul><li><a href="/wiki/GNGT1" title="GNGT1">GNGT1</a></li> <li><a href="/wiki/GNGT2" title="GNGT2">GNGT2</a></li> <li><a href="/wiki/GNG2" title="GNG2">GNG2</a></li> <li><a href="/wiki/GNG3" title="GNG3">GNG3</a></li> <li><a href="/wiki/GNG4" title="GNG4">GNG4</a></li> <li><a href="/wiki/GNG5" title="GNG5">GNG5</a></li> <li><a href="/wiki/GNG7" title="GNG7">GNG7</a></li> <li><a href="/w/index.php?title=GNG8&action=edit&redlink=1" class="new" title="GNG8 (page does not exist)">GNG8</a></li> <li><a href="/w/index.php?title=GNG10&action=edit&redlink=1" class="new" title="GNG10 (page does not exist)">GNG10</a></li> <li><a href="/wiki/GNG11" title="GNG11">GNG11</a></li> <li><a href="/wiki/GNG12" title="GNG12">GNG12</a></li> <li><a href="/wiki/GNG13" title="GNG13">GNG13</a></li> <li><a href="/wiki/BSCL2" title="BSCL2">BSCL2</a></li></ul></li></ul> </div></td></tr><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/G_protein-coupled_receptor_kinase" title="G protein-coupled receptor kinase">G protein-coupled receptor kinase</a></li> <li><a href="/wiki/AMP-activated_protein_kinase" title="AMP-activated protein kinase">AMP-activated protein kinase</a></li></ul> </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/Small_GTPase" title="Small GTPase">Monomeric</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/ADP_ribosylation_factor" title="ADP ribosylation factor">ARFs</a></li> <li><a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rabs</a></li> <li><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/Rho_family_of_GTPases" title="Rho family of GTPases">Rhos</a></li> <li><a href="/wiki/ADP_ribosylation_factor" title="ADP ribosylation factor">Arfs</a></li> <li><a href="/wiki/Ran_(gene)" class="mw-redirect" title="Ran (gene)">Ran</a></li> <li><a href="/wiki/RHEB" title="RHEB">Rhebs</a></li> <li><a href="/wiki/Rap_GTP-binding_protein" title="Rap GTP-binding protein">Raps</a></li> <li><a href="/wiki/RRAD_(gene)" title="RRAD (gene)">RGKs</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Cyclin</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/Cyclin-dependent_kinase_inhibitor_protein" title="Cyclin-dependent kinase inhibitor protein">Cyclin-dependent kinase inhibitor protein</a></li> <li><a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">Cyclin-dependent kinase</a></li> <li><a href="/wiki/Cyclin" title="Cyclin">Cyclin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Lipid</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/Phosphoinositide_phospholipase_C" title="Phosphoinositide phospholipase C">Phosphoinositide phospholipase C</a></li> <li><a href="/wiki/Phospholipase_C" title="Phospholipase C">Phospholipase C</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Other <a href="/wiki/Protein_kinase" title="Protein kinase">protein kinase</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/Serine/threonine-specific_protein_kinase" title="Serine/threonine-specific protein kinase">Serine/threonine</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/Casein_kinase" title="Casein kinase">Casein kinase</a> <ul><li><a href="/wiki/Casein_kinase_1" title="Casein kinase 1">1</a></li> <li><a href="/wiki/Casein_kinase_2" title="Casein kinase 2">2</a></li></ul></li> <li><a href="/wiki/EIF-2_kinase" title="EIF-2 kinase">eIF-2 kinase</a> <ul><li><a href="/wiki/EIF2AK3" title="EIF2AK3">EIF2AK3</a></li></ul></li> <li><a href="/wiki/Glycogen_synthase_kinase" class="mw-redirect" title="Glycogen synthase kinase">Glycogen synthase kinase</a> <ul><li><a href="/w/index.php?title=GSK1&action=edit&redlink=1" class="new" title="GSK1 (page does not exist)">GSK1</a></li> <li><a href="/w/index.php?title=GSK2&action=edit&redlink=1" class="new" title="GSK2 (page does not exist)">GSK2</a></li> <li><a href="/wiki/GSK-3" title="GSK-3">GSK-3</a></li> <li><a href="/wiki/GSK3A" title="GSK3A">GSK3A</a></li> <li><a href="/wiki/GSK3B" class="mw-redirect" title="GSK3B">GSK3B</a></li></ul></li> <li><a href="/wiki/I%CE%BAB_kinase" title="IκB kinase">IκB kinase</a> <ul><li><a href="/wiki/CHUK" title="CHUK">CHUK</a></li> <li><a href="/wiki/IKK2" title="IKK2">IKK2</a></li> <li><a href="/wiki/IKBKG" title="IKBKG">IKBKG</a></li></ul></li> <li><a href="/wiki/Interleukin-1_receptor_associated_kinase" title="Interleukin-1 receptor associated kinase">Interleukin-1 receptor associated kinase</a> <ul><li><a href="/wiki/IRAK1" title="IRAK1">IRAK1</a></li> <li><a href="/wiki/IRAK2" title="IRAK2">IRAK2</a></li> <li><a href="/wiki/IRAK3" title="IRAK3">IRAK3</a></li> <li><a href="/wiki/IRAK4" title="IRAK4">IRAK4</a></li></ul></li> <li><a href="/wiki/Lim_kinase" title="Lim kinase">Lim kinase</a> <ul><li><a href="/wiki/LIMK1" title="LIMK1">LIMK1</a></li> <li><a href="/wiki/LIMK2" title="LIMK2">LIMK2</a></li></ul></li> <li><a href="/wiki/P21-activated_kinases" title="P21-activated kinases">p21-activated kinases</a> <ul><li><a href="/wiki/PAK1" title="PAK1">PAK1</a></li> <li><a href="/wiki/PAK2" title="PAK2">PAK2</a></li> <li><a href="/wiki/PAK3" title="PAK3">PAK3</a></li> <li><a href="/wiki/PAK4" title="PAK4">PAK4</a></li></ul></li> <li><a href="/wiki/Rho-associated_protein_kinase" title="Rho-associated protein kinase">Rho-associated protein kinase</a> <ul><li><a href="/wiki/ROCK1" title="ROCK1">ROCK1</a></li> <li><a href="/wiki/ROCK2" title="ROCK2">ROCK2</a></li></ul></li> <li><a href="/wiki/Ribosomal_s6_kinase" title="Ribosomal s6 kinase">Ribosomal s6 kinase</a> <ul><li><a href="/wiki/RPS6KA1" title="RPS6KA1">RPS6KA1</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Non-receptor_tyrosine_kinase" title="Non-receptor tyrosine kinase">Tyrosine</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/ZAP70" title="ZAP70">ZAP70</a></li> <li><a href="/w/index.php?title=Focal_adhesion_protein-tyrosine_kinase&action=edit&redlink=1" class="new" title="Focal adhesion protein-tyrosine kinase (page does not exist)">Focal adhesion protein-tyrosine kinase</a> <ul><li><a href="/wiki/PTK2" title="PTK2">PTK2</a></li> <li><a href="/wiki/PTK2B" title="PTK2B">PTK2B</a></li></ul></li> <li><a href="/wiki/Bruton%27s_tyrosine_kinase" title="Bruton's tyrosine kinase">BTK</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Serine/threonine/tyrosine</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/Dual-specificity_kinase" title="Dual-specificity kinase">Dual-specificity kinase</a> <ul><li><a href="/wiki/DYRK1A" title="DYRK1A">DYRK1A</a></li> <li><a href="/wiki/DYRK1B" title="DYRK1B">DYRK1B</a></li> <li><a href="/wiki/DYRK2" title="DYRK2">DYRK2</a></li> <li><a href="/wiki/DYRK3" title="DYRK3">DYRK3</a></li> <li><a href="/w/index.php?title=DYRK4&action=edit&redlink=1" class="new" title="DYRK4 (page does not exist)">DYRK4</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Arginine</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/Arginine_kinase" title="Arginine kinase">Arginine kinase</a></li> <li><a href="/w/index.php?title=McsB&action=edit&redlink=1" class="new" title="McsB (page does not exist)">McsB</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Other <a href="/wiki/Protein_phosphatase" title="Protein phosphatase">protein phosphatase</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/Protein_serine/threonine_phosphatase" title="Protein serine/threonine phosphatase">Serine/threonine</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/Protein_phosphatase_2" title="Protein phosphatase 2">Protein phosphatase 2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Protein_tyrosine_phosphatase" title="Protein tyrosine phosphatase">Tyrosine</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/Protein_tyrosine_phosphatase" title="Protein tyrosine phosphatase">protein tyrosine phosphatase</a>: <a href="/w/index.php?title=Receptor-like_protein_tyrosine_phosphatase&action=edit&redlink=1" class="new" title="Receptor-like protein tyrosine phosphatase (page does not exist)">Receptor-like protein tyrosine phosphatase</a></li> <li><a href="/wiki/Sh2_domain-containing_protein_tyrosine_phosphatase" class="mw-redirect" title="Sh2 domain-containing protein tyrosine phosphatase">Sh2 domain-containing protein tyrosine phosphatase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">both:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dual-specificity_phosphatase" title="Dual-specificity phosphatase">Dual-specificity phosphatase</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%"><a href="/wiki/Apoptosis" title="Apoptosis">Apoptosis</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><i>see <a href="/wiki/Template:Apoptosis_signaling_pathway" title="Template:Apoptosis signaling pathway">apoptosis signaling pathway</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%"><a href="/wiki/GTP-binding_protein_regulators" title="GTP-binding protein regulators">GTP-binding protein regulators</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><i>see <a href="/wiki/Template:GTP-binding_protein_regulators" title="Template:GTP-binding protein regulators">GTP-binding protein regulators</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background-color: AntiqueWhite;width:1%">Other</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/ATF6" title="ATF6">Activating transcription factor 6</a></li> <li><a href="/wiki/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">Signal transducing adaptor protein</a></li> <li><a href="/wiki/I%CE%BAB%CE%B1" title="IκBα">I-kappa B protein</a></li> <li><a href="/wiki/MUC4" class="mw-redirect" title="MUC4">Mucin-4</a></li> <li><a href="/wiki/Olfactory_marker_protein" title="Olfactory marker protein">Olfactory marker protein</a></li> <li><a href="/wiki/Phosphatidylethanolamine_binding_protein" class="mw-redirect" title="Phosphatidylethanolamine binding protein">Phosphatidylethanolamine binding protein</a></li> <li><a href="/wiki/EDARADD" title="EDARADD">EDARADD</a></li> <li><a href="/wiki/PRKCSH" title="PRKCSH">PRKCSH</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:Deficiencies_of_intracellular_signaling_peptides_and_proteins" title="Template:Deficiencies of intracellular signaling peptides and proteins">deficiencies of intracellular signaling peptides and proteins</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="Tumor_suppressor_genes_and_Oncogenes" 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:Tumor_suppressor_genes_and_oncogenes" title="Template:Tumor suppressor genes and oncogenes"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Tumor_suppressor_genes_and_oncogenes" title="Template talk:Tumor suppressor genes and oncogenes"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Tumor_suppressor_genes_and_oncogenes" title="Special:EditPage/Template:Tumor suppressor genes and oncogenes"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Tumor_suppressor_genes_and_Oncogenes" style="font-size:114%;margin:0 4em"><a href="/wiki/Tumor_suppressor_gene" title="Tumor suppressor gene">Tumor suppressor genes</a> and <a href="/wiki/Oncogene" title="Oncogene">Oncogenes</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</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 id="Growth_factors" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Growth_factor" title="Growth factor">Growth factors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</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/Platelet-derived_growth_factor" title="Platelet-derived growth factor">c-Sis/PDGF</a></li> <li><a href="/wiki/Hepatocyte_growth_factor" title="Hepatocyte growth factor">HGF</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptor</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%"><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/CDH1_(gene)" class="mw-redirect" title="CDH1 (gene)">CDH1</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/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/PTCH1" title="PTCH1">PTCH1</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/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/TGF_beta_receptor_2" title="TGF beta receptor 2">TGF beta receptor 2</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/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">Receptor tyrosine kinase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</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/ErbB" title="ErbB">ErbB</a>/<a href="/wiki/Epidermal_growth_factor_receptor" title="Epidermal growth factor receptor">c-ErbB</a> <ul><li><a href="/wiki/HER2/neu" class="mw-redirect" title="HER2/neu">HER2/neu</a></li> <li><a href="/wiki/ERBB3" title="ERBB3">Her 3</a></li></ul></li> <li><a href="/wiki/C-Met" class="mw-redirect" title="C-Met">c-Met</a></li> <li><a href="/wiki/RET_proto-oncogene" title="RET proto-oncogene">c-Ret</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/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="ONCO" scope="row" class="navbox-group" style="width:1%">ONCO</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/CD117" class="mw-redirect" title="CD117">c-Kit</a></li> <li><a href="/wiki/CD135" title="CD135">Flt3</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular signaling P+Ps</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%"><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/Beta-catenin" class="mw-redirect" title="Beta-catenin">Beta-catenin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/Adenomatosis_polyposis_coli" class="mw-redirect" title="Adenomatosis polyposis coli">APC</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/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/Mothers_against_decapentaplegic_homolog_2" title="Mothers against decapentaplegic homolog 2">SMAD2</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_4" title="Mothers against decapentaplegic homolog 4">SMAD4</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/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/AKT" class="mw-redirect" title="AKT">c-Akt</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/PTEN_(gene)" title="PTEN (gene)">PTEN</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/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo signaling pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/Merlin_(protein)" title="Merlin (protein)">Neurofibromin 2/Merlin</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/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK pathway</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/Ras_(protein)" class="mw-redirect" title="Ras (protein)">c-Ras</a></li> <li><a href="/wiki/HRAS" title="HRAS">HRAS</a></li> <li><a href="/wiki/C-Raf" title="C-Raf">c-Raf</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/Neurofibromin" title="Neurofibromin">Neurofibromin 1</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/unknown</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%">ONCO</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/Proto-oncogene_tyrosine-protein_kinase_Src" title="Proto-oncogene tyrosine-protein kinase Src">c-Src</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/Maspin" title="Maspin">Maspin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</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%"><a href="/wiki/Cell_cycle" title="Cell cycle">Cell cycle</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4</a></li> <li><a href="/wiki/Cyclin_D" title="Cyclin D">Cyclin D</a></li> <li><a href="/wiki/Cyclin_E" title="Cyclin E">Cyclin E</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/P53" title="P53">p53</a></li> <li><a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">pRb</a></li> <li><a href="/wiki/WT1" class="mw-redirect" title="WT1">WT1</a></li> <li><a href="/wiki/P16_(gene)" class="mw-redirect" title="P16 (gene)">p16</a>/<a href="/wiki/P14arf" title="P14arf">p14arf</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/DNA_repair" title="DNA repair">DNA repair</a>/<a href="/wiki/Fanconi_anemia" title="Fanconi anemia">Fanconi</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TSP" scope="row" class="navbox-group" style="width:1%">TSP</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/BRCA1" title="BRCA1">BRCA1</a></li> <li><a href="/wiki/BRCA2" title="BRCA2">BRCA2</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/Ubiquitin_ligase" title="Ubiquitin ligase">Ubiquitin ligase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/CBL_(gene)" title="CBL (gene)">CBL</a></li> <li><a href="/wiki/Mdm2" title="Mdm2">MDM2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/Von_Hippel%E2%80%93Lindau_tumor_suppressor" title="Von Hippel–Lindau tumor suppressor">VHL</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/Transcription_factor" title="Transcription factor">Transcription factor</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">ONCO</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/AP-1_transcription_factor" title="AP-1 transcription factor">AP-1</a> <ul><li><a href="/wiki/C-Fos" class="mw-redirect" title="C-Fos">c-Fos</a></li> <li><a href="/wiki/C-jun" class="mw-redirect" title="C-jun">c-Jun</a></li></ul></li> <li><a href="/wiki/Myc" title="Myc">c-Myc</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">TSP</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/KLF6" title="KLF6">KLF6</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</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 id="Apoptosis_inhibitor" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">Apoptosis inhibitor</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/SDHB" title="SDHB">SDHB</a></li> <li><a href="/wiki/SDHD" title="SDHD">SDHD</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/ungrouped</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/Bcl-2" title="Bcl-2">c-Bcl-2</a></li> <li><a href="/wiki/Notch_signaling" class="mw-redirect" title="Notch signaling">Notch</a></li> <li><a href="/wiki/Stathmin" title="Stathmin">Stathmin</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="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" 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 .portal-bar-bordered{padding:0 2em;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em auto 0}.mw-parser-output .portal-bar-related{font-size:100%;justify-content:flex-start}.mw-parser-output .portal-bar-unbordered{padding:0 1.7em;margin-left:0}.mw-parser-output .portal-bar-header{margin:0 1em 0 0.5em;flex:0 0 auto;min-height:24px}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;padding:0.15em 0;column-gap:1em;align-items:baseline;margin:0;list-style:none}.mw-parser-output .portal-bar-content-related{margin:0;list-style:none}.mw-parser-output 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