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class="vector-toc-numb">2</span> <span>Classification</span> </div> </a> <ul id="toc-Classification-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Physiological_roles" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Physiological_roles"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Physiological roles</span> </div> </a> <ul id="toc-Physiological_roles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Receptor_structure" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Receptor_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Receptor structure</span> </div> </a> <ul id="toc-Receptor_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structure–function_relationships" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Structure–function_relationships"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Structure–function relationships</span> </div> </a> <ul id="toc-Structure–function_relationships-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanism" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mechanism"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Mechanism</span> </div> </a> <button aria-controls="toc-Mechanism-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 Mechanism subsection</span> </button> <ul id="toc-Mechanism-sublist" class="vector-toc-list"> <li id="toc-Ligand_binding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ligand_binding"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Ligand binding</span> </div> </a> <ul id="toc-Ligand_binding-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Conformational_change" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Conformational_change"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Conformational change</span> </div> </a> <ul id="toc-Conformational_change-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-G-protein_activation/deactivation_cycle" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#G-protein_activation/deactivation_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>G-protein activation/deactivation cycle</span> </div> </a> <ul id="toc-G-protein_activation/deactivation_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Crosstalk" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Crosstalk"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span>Crosstalk</span> </div> </a> <ul id="toc-Crosstalk-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Signaling" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Signaling</span> </div> </a> <button aria-controls="toc-Signaling-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 Signaling subsection</span> </button> <ul id="toc-Signaling-sublist" class="vector-toc-list"> <li id="toc-G-protein-dependent_signaling" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#G-protein-dependent_signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>G-protein-dependent signaling</span> </div> </a> <ul id="toc-G-protein-dependent_signaling-sublist" class="vector-toc-list"> <li id="toc-Gα_signaling" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Gα_signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.1</span> <span>Gα signaling</span> </div> </a> <ul id="toc-Gα_signaling-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gβγ_signaling" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Gβγ_signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.2</span> <span>Gβγ signaling</span> </div> </a> <ul id="toc-Gβγ_signaling-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-G-protein-independent_signaling" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#G-protein-independent_signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>G-protein-independent signaling</span> </div> </a> <ul id="toc-G-protein-independent_signaling-sublist" class="vector-toc-list"> <li id="toc-Examples" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Examples"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2.1</span> <span>Examples</span> </div> </a> <ul id="toc-Examples-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-GPCR-independent_signaling_by_heterotrimeric_G-proteins" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#GPCR-independent_signaling_by_heterotrimeric_G-proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2.2</span> <span>GPCR-independent signaling by heterotrimeric G-proteins</span> </div> </a> <ul id="toc-GPCR-independent_signaling_by_heterotrimeric_G-proteins-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Details_of_cAMP_and_PIP2_pathways" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Details_of_cAMP_and_PIP2_pathways"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Details of cAMP and PIP2 pathways</span> </div> </a> <button aria-controls="toc-Details_of_cAMP_and_PIP2_pathways-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 Details of cAMP and PIP2 pathways subsection</span> </button> <ul id="toc-Details_of_cAMP_and_PIP2_pathways-sublist" class="vector-toc-list"> <li id="toc-cAMP_signal_pathway" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#cAMP_signal_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>cAMP signal pathway</span> </div> </a> <ul id="toc-cAMP_signal_pathway-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phosphatidylinositol_signal_pathway" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phosphatidylinositol_signal_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Phosphatidylinositol signal pathway</span> </div> </a> <ul id="toc-Phosphatidylinositol_signal_pathway-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Receptor_regulation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Receptor_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Receptor regulation</span> </div> </a> <button aria-controls="toc-Receptor_regulation-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 Receptor regulation subsection</span> </button> <ul id="toc-Receptor_regulation-sublist" class="vector-toc-list"> <li id="toc-Phosphorylation_by_cAMP-dependent_protein_kinases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phosphorylation_by_cAMP-dependent_protein_kinases"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.1</span> <span>Phosphorylation by cAMP-dependent protein kinases</span> </div> </a> <ul id="toc-Phosphorylation_by_cAMP-dependent_protein_kinases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phosphorylation_by_GRKs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phosphorylation_by_GRKs"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.2</span> <span>Phosphorylation by GRKs</span> </div> </a> <ul id="toc-Phosphorylation_by_GRKs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanisms_of_GPCR_signal_termination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mechanisms_of_GPCR_signal_termination"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.3</span> <span>Mechanisms of GPCR signal termination</span> </div> </a> <ul id="toc-Mechanisms_of_GPCR_signal_termination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-GPCR_cellular_regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#GPCR_cellular_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.4</span> <span>GPCR cellular regulation</span> </div> </a> <ul id="toc-GPCR_cellular_regulation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Receptor_oligomerization" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Receptor_oligomerization"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Receptor oligomerization</span> </div> </a> <ul id="toc-Receptor_oligomerization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Origin_and_diversification_of_the_superfamily" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Origin_and_diversification_of_the_superfamily"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Origin and diversification of the superfamily</span> </div> </a> <ul id="toc-Origin_and_diversification_of_the_superfamily-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</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"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">14</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"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">15</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">G protein-coupled receptor</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 42 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-42" 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">42 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%85%D8%B3%D8%AA%D9%82%D8%A8%D9%84_%D9%85%D9%82%D8%AA%D8%B1%D9%86_%D8%A8%D8%A7%D9%84%D8%A8%D8%B1%D9%88%D8%AA%D9%8A%D9%86_%D8%AC" title="مستقبل مقترن بالبروتين ج – Arabic" lang="ar" hreflang="ar" data-title="مستقبل مقترن بالبروتين ج" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Receptori_spregnuti_sa_G-proteinom" title="Receptori spregnuti sa G-proteinom – Bosnian" lang="bs" hreflang="bs" data-title="Receptori spregnuti sa G-proteinom" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Receptor_acoblat_a_prote%C3%AFnes_G" title="Receptor acoblat a proteïnes G – Catalan" lang="ca" hreflang="ca" data-title="Receptor acoblat a proteïnes G" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Receptor_sp%C5%99a%C5%BEen%C3%BD_s_G_proteinem" title="Receptor spřažený s G proteinem – Czech" lang="cs" hreflang="cs" data-title="Receptor spřažený s G proteinem" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/G-protein-koblede_receptorer" title="G-protein-koblede receptorer – Danish" lang="da" hreflang="da" data-title="G-protein-koblede receptorer" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://de.wikipedia.org/wiki/G-Protein-gekoppelte_Rezeptoren" title="G-Protein-gekoppelte Rezeptoren – German" lang="de" hreflang="de" data-title="G-Protein-gekoppelte Rezeptoren" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/G-valguga_seotud_retseptorid" title="G-valguga seotud retseptorid – Estonian" lang="et" hreflang="et" data-title="G-valguga seotud retseptorid" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A5%CF%80%CE%BF%CE%B4%CE%BF%CF%87%CE%AD%CE%B1%CF%82_%CF%83%CF%85%CE%B6%CE%B5%CF%85%CE%B3%CE%BC%CE%AD%CE%BD%CE%BF%CF%82_%CE%BC%CE%B5_%CF%80%CF%81%CF%89%CF%84%CE%B5%CE%90%CE%BD%CE%B7_G" title="Υποδοχέας συζευγμένος με πρωτεΐνη G – Greek" lang="el" hreflang="el" data-title="Υποδοχέας συζευγμένος με πρωτεΐνη G" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Receptor_acoplado_a_prote%C3%ADnas_G" title="Receptor acoplado a proteínas G – Spanish" lang="es" hreflang="es" data-title="Receptor acoplado a proteínas G" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/G_proteinei_loturiko_hartzaile" title="G proteinei loturiko hartzaile – Basque" lang="eu" hreflang="eu" data-title="G proteinei loturiko hartzaile" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%AF%DB%8C%D8%B1%D9%86%D8%AF%D9%87_%D8%AC%D9%81%D8%AA%E2%80%8C%D8%B4%D9%88%D9%86%D8%AF%D9%87_%D8%A8%D8%A7_%D9%BE%D8%B1%D9%88%D8%AA%D8%A6%DB%8C%D9%86_%D8%AC%DB%8C" title="گیرنده جفت‌شونده با پروتئین جی – Persian" lang="fa" hreflang="fa" data-title="گیرنده جفت‌شونده با پروتئین جی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/R%C3%A9cepteur_coupl%C3%A9_aux_prot%C3%A9ines_G" title="Récepteur couplé aux protéines G – French" lang="fr" hreflang="fr" data-title="Récepteur couplé aux protéines G" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Receptor_acoplado_%C3%A1_prote%C3%ADna_G" title="Receptor acoplado á proteína G – Galician" lang="gl" hreflang="gl" data-title="Receptor acoplado á proteína G" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/G_%EB%8B%A8%EB%B0%B1%EC%A7%88_%EC%97%B0%EA%B2%B0_%EC%88%98%EC%9A%A9%EC%B2%B4" title="G 단백질 연결 수용체 – Korean" lang="ko" hreflang="ko" data-title="G 단백질 연결 수용체" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Reseptor_terhubung_protein_G" title="Reseptor terhubung protein G – Indonesian" lang="id" hreflang="id" data-title="Reseptor terhubung protein G" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Recettori_accoppiati_a_proteine_G" title="Recettori accoppiati a proteine G – Italian" lang="it" hreflang="it" data-title="Recettori accoppiati a proteine G" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A7%D7%95%D7%9C%D7%98%D7%9F_%D7%94%D7%9E%D7%A6%D7%95%D7%9E%D7%93_%D7%9C%D7%97%D7%9C%D7%91%D7%95%D7%9F_G" title="קולטן המצומד לחלבון G – Hebrew" lang="he" hreflang="he" data-title="קולטן המצומד לחלבון G" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/G-%E1%83%AA%E1%83%98%E1%83%9A%E1%83%90%E1%83%A1%E1%83%97%E1%83%90%E1%83%9C_%E1%83%93%E1%83%90%E1%83%99%E1%83%90%E1%83%95%E1%83%A8%E1%83%98%E1%83%A0%E1%83%94%E1%83%91%E1%83%A3%E1%83%9A%E1%83%98_%E1%83%A0%E1%83%94%E1%83%AA%E1%83%94%E1%83%9E%E1%83%A2%E1%83%9D%E1%83%A0%E1%83%98" title="G-ცილასთან დაკავშირებული რეცეპტორი – Georgian" lang="ka" hreflang="ka" data-title="G-ცილასთან დაკავშირებული რეცეპტორი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Receptorium_proteino_G_copulatum" title="Receptorium proteino G copulatum – Latin" lang="la" hreflang="la" data-title="Receptorium proteino G copulatum" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Ar_G_prote%C4%ABniem_saist%C4%ABtie_receptori" title="Ar G proteīniem saistītie receptori – Latvian" lang="lv" hreflang="lv" data-title="Ar G proteīniem saistītie receptori" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/G-protein-kapcsolt_receptor" title="G-protein-kapcsolt receptor – Hungarian" lang="hu" hreflang="hu" data-title="G-protein-kapcsolt receptor" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Reseptor_bergandingan_protein_G" title="Reseptor bergandingan protein G – Malay" lang="ms" hreflang="ms" data-title="Reseptor bergandingan protein G" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/G-prote%C3%AFnegekoppelde_receptor" title="G-proteïnegekoppelde receptor – Dutch" lang="nl" hreflang="nl" data-title="G-proteïnegekoppelde receptor" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-new mw-list-item"><a href="https://new.wikipedia.org/wiki/%E0%A4%9C%E0%A4%BF_%E0%A4%AA%E0%A5%8D%E0%A4%B0%E0%A5%8B%E0%A4%9F%E0%A4%BF%E0%A4%A8_%E0%A4%95%E0%A4%AA%E0%A4%B2%E0%A5%8D%E0%A4%A1_%E0%A4%B0%E0%A4%BF%E0%A4%B8%E0%A5%87%E0%A4%AA%E0%A5%8D%E0%A4%9F%E0%A4%B0" title="जि प्रोटिन कपल्ड रिसेप्टर – Newari" lang="new" hreflang="new" data-title="जि प्रोटिन कपल्ड रिसेप्टर" data-language-autonym="नेपाल भाषा" data-language-local-name="Newari" class="interlanguage-link-target"><span>नेपाल भाषा</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/G%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA%E5%85%B1%E5%BD%B9%E5%8F%97%E5%AE%B9%E4%BD%93" title="Gタンパク質共役受容体 – Japanese" lang="ja" hreflang="ja" data-title="Gタンパク質共役受容体" 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-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/G-protein-kopla_reseptor" title="G-protein-kopla reseptor – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="G-protein-kopla reseptor" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/GPCR_retseptorlar" title="GPCR retseptorlar – Uzbek" lang="uz" hreflang="uz" data-title="GPCR retseptorlar" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Receptory_sprz%C4%99%C5%BCone_z_bia%C5%82kami_G" title="Receptory sprzężone z białkami G – Polish" lang="pl" hreflang="pl" data-title="Receptory sprzężone z białkami G" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Receptor_acoplado_%C3%A0_prote%C3%ADna_G" title="Receptor acoplado à proteína G – Portuguese" lang="pt" hreflang="pt" data-title="Receptor acoplado à proteína G" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Receptor_cuplat_cu_proteina_G" title="Receptor cuplat cu proteina G – Romanian" lang="ro" hreflang="ro" data-title="Receptor cuplat cu proteina G" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/GPCR" title="GPCR – Russian" lang="ru" hreflang="ru" data-title="GPCR" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Receptor_spriahnut%C3%BD_s_G-prote%C3%ADnmi" title="Receptor spriahnutý s G-proteínmi – Slovak" lang="sk" hreflang="sk" data-title="Receptor spriahnutý s G-proteínmi" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sr badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://sr.wikipedia.org/wiki/Receptori_spregnuti_s_G_proteinom" title="Receptori spregnuti s G proteinom – Serbian" lang="sr" hreflang="sr" data-title="Receptori spregnuti s G proteinom" 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/G_protein_spregnuti_receptori" title="G protein spregnuti receptori – Serbo-Croatian" lang="sh" hreflang="sh" data-title="G protein spregnuti receptori" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/G-proteiinikytkent%C3%A4iset_reseptorit" title="G-proteiinikytkentäiset reseptorit – Finnish" lang="fi" hreflang="fi" data-title="G-proteiinikytkentäiset reseptorit" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/G-proteinkopplade_receptorer" title="G-proteinkopplade receptorer – Swedish" lang="sv" hreflang="sv" data-title="G-proteinkopplade receptorer" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%AB%E0%B8%99%E0%B9%88%E0%B8%A7%E0%B8%A2%E0%B8%A3%E0%B8%B1%E0%B8%9A%E0%B8%97%E0%B8%B5%E0%B9%88%E0%B8%88%E0%B8%B1%E0%B8%9A%E0%B8%84%E0%B8%B9%E0%B9%88%E0%B8%81%E0%B8%B1%E0%B8%9A%E0%B8%88%E0%B8%B5%E0%B9%82%E0%B8%9B%E0%B8%A3%E0%B8%95%E0%B8%B5%E0%B8%99" title="หน่วยรับที่จับคู่กับจีโปรตีน – Thai" lang="th" hreflang="th" data-title="หน่วยรับที่จับคู่กับจีโปรตีน" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/G_proteini_kenetli_resept%C3%B6r" title="G proteini kenetli reseptör – Turkish" lang="tr" hreflang="tr" data-title="G proteini kenetli reseptör" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/G-%D0%B1%D1%96%D0%BB%D0%BE%D0%BA%D1%81%D0%BF%D1%80%D1%8F%D0%B6%D0%B5%D0%BD%D1%96_%D1%80%D0%B5%D1%86%D0%B5%D0%BF%D1%82%D0%BE%D1%80%D0%B8" title="G-білокспряжені рецептори – Ukrainian" lang="uk" hreflang="uk" data-title="G-білокспряжені рецептори" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Th%E1%BB%A5_th%E1%BB%83_b%E1%BA%AFt_c%E1%BA%B7p_v%E1%BB%9Bi_G_protein" title="Thụ thể bắt cặp với G protein – Vietnamese" lang="vi" hreflang="vi" data-title="Thụ thể bắt cặp với G protein" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a 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dir="ltr"><style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">"GPCR" redirects here. For the Great Proletarian Cultural Revolution, see <a href="/wiki/Cultural_Revolution" title="Cultural Revolution">Cultural Revolution</a>.</div> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Class of cell surface receptors coupled to G-protein-associated intracellular signaling</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">GPCR</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Beta-2-adrenergic-receptor.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Beta-2-adrenergic-receptor.png/220px-Beta-2-adrenergic-receptor.png" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Beta-2-adrenergic-receptor.png/330px-Beta-2-adrenergic-receptor.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Beta-2-adrenergic-receptor.png/440px-Beta-2-adrenergic-receptor.png 2x" data-file-width="768" data-file-height="1024" /></a></span><div class="infobox-caption">The human <a href="/wiki/Beta-2_adrenergic_receptor" title="Beta-2 adrenergic receptor">beta-2 adrenergic receptor</a> in complex with the partial <a href="/wiki/Inverse_agonist" title="Inverse agonist">inverse agonist</a> <a href="/wiki/Carazolol" title="Carazolol">carazolol</a><sup id="cite_ref-Cherezov_2007_1-0" class="reference"><a href="#cite_note-Cherezov_2007-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></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">7tm_1</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/PF00001">PF00001</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Pfam" title="Pfam">Pfam</a> clan</th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/set/pfam/CL0192/">CL0192</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/ECOD" class="mw-redirect" title="ECOD">ECOD</a></th><td class="infobox-data ECOD" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/tree?id=5001.1.1">5001.1.1</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/IPR000276">IPR000276</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/PDOC00210">PDOC00210</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/TCDB" class="mw-redirect" title="TCDB">TCDB</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://www.tcdb.org/search/result.php?tc=9.A.14">9.A.14</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM superfamily</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/protein_superfamilies/6">6</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM protein</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/proteins?search=1gzm">1gzm</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=cd14964">cd14964</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/PF00001?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00001">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:PF00001%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:PF00001">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF00001">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=PF00001">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:PDB_1hzx_7TM_Sketch_Membrane.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PDB_1hzx_7TM_Sketch_Membrane.png/220px-PDB_1hzx_7TM_Sketch_Membrane.png" decoding="async" width="220" height="154" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PDB_1hzx_7TM_Sketch_Membrane.png/330px-PDB_1hzx_7TM_Sketch_Membrane.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PDB_1hzx_7TM_Sketch_Membrane.png/440px-PDB_1hzx_7TM_Sketch_Membrane.png 2x" data-file-width="1029" data-file-height="720" /></a><figcaption>The seven-transmembrane α-helix structure of <a href="/wiki/Bovine" class="mw-redirect" title="Bovine">bovine</a> rhodopsin</figcaption></figure> <p><b>G protein-coupled receptors</b> (<b>GPCRs</b>), also known as <b>seven-(pass)-transmembrane domain receptors</b>, <b>7TM receptors</b>, <b>heptahelical receptors</b>, <b>serpentine receptors</b>, and <b>G protein-linked receptors</b> (<b>GPLR</b>), form a large <a href="/wiki/Protein_family" title="Protein family">group of evolutionarily related proteins</a> that are <a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">cell surface receptors</a> that detect <a href="/wiki/Molecule" title="Molecule">molecules</a> outside the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> and activate cellular responses. They are coupled with <a href="/wiki/G_protein" title="G protein">G proteins</a>. They pass through the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> seven times in the form of six loops<sup id="cite_ref-Zhang2013_2-0" class="reference"><a href="#cite_note-Zhang2013-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> (three extracellular loops interacting with <a href="/wiki/Ligand" title="Ligand">ligand</a> molecules, three intracellular loops interacting with G proteins, an N-terminal extracellular region and a C-terminal intracellular region<sup id="cite_ref-Zhang2013_2-1" class="reference"><a href="#cite_note-Zhang2013-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>) of <a href="/wiki/Amino_acid_residue" class="mw-redirect" title="Amino acid residue">amino acid residues</a>, which is why they are sometimes referred to as seven-transmembrane receptors.<sup id="cite_ref-Trzaskowski2012_3-0" class="reference"><a href="#cite_note-Trzaskowski2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Ligands can bind either to the extracellular N-terminus and loops (e.g. glutamate receptors) or to the binding site within transmembrane helices (<a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a>-like family). They are all activated by <a href="/wiki/Agonist" title="Agonist">agonists</a>, although a spontaneous auto-activation of an empty receptor has also been observed.<sup id="cite_ref-Trzaskowski2012_3-1" class="reference"><a href="#cite_note-Trzaskowski2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>G protein-coupled receptors are found only in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, including <a href="/wiki/Yeast" title="Yeast">yeast</a>, and <a href="/wiki/Choanoflagellate" title="Choanoflagellate">choanoflagellates</a>.<sup id="cite_ref-pmid12869759_4-0" class="reference"><a href="#cite_note-pmid12869759-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligands</a> that bind and activate these receptors include light-sensitive compounds, <a href="/wiki/Odor" title="Odor">odors</a>, <a href="/wiki/Pheromone" title="Pheromone">pheromones</a>, <a href="/wiki/Hormone" title="Hormone">hormones</a>, and <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitters</a>, and vary in size from small molecules to <a href="/wiki/Peptide" title="Peptide">peptides</a> to large <a href="/wiki/Protein" title="Protein">proteins</a>. G protein-coupled receptors are involved in many diseases. </p><p>There are two principal signal transduction pathways involving the G protein-coupled receptors: </p> <ul><li>the <a href="/wiki/Cyclic_adenosine_monophosphate" title="Cyclic adenosine monophosphate">cAMP</a> signal pathway and</li> <li>the <a href="/wiki/Phosphatidylinositol" title="Phosphatidylinositol">phosphatidylinositol</a> signal pathway.<sup id="cite_ref-Gilman_1987_5-0" class="reference"><a href="#cite_note-Gilman_1987-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup></li></ul> <p>When a ligand binds to the GPCR it causes a conformational change in the GPCR, which allows it to act as a <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">guanine nucleotide exchange factor</a> (GEF). The GPCR can then activate an associated <a href="/wiki/G_protein" title="G protein">G protein</a> by exchanging the <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">GDP</a> bound to the G protein for a <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a>. The G protein's α subunit, together with the bound GTP, can then dissociate from the β and γ subunits to further affect intracellular signaling proteins or target functional proteins directly depending on the α subunit type (<a href="/wiki/Gs_alpha_subunit" title="Gs alpha subunit">G<sub>αs</sub></a>, <a href="/wiki/Gi_alpha_subunit" title="Gi alpha subunit">G<sub>αi/o</sub></a>, <a href="/wiki/Gq_alpha_subunit" title="Gq alpha subunit">G<sub>αq/11</sub></a>, <a href="/wiki/G12/G13_alpha_subunits" title="G12/G13 alpha subunits">G<sub>α12/13</sub></a>).<sup id="cite_ref-Wettschureck_2005_6-0" class="reference"><a href="#cite_note-Wettschureck_2005-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 1160">: 1160 </span></sup> </p><p>GPCRs are an important drug target and approximately 34%<sup id="cite_ref-Hauser_2018_7-0" class="reference"><a href="#cite_note-Hauser_2018-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> of all Food and Drug Administration (FDA) approved drugs target 108 members of this family. The global sales volume for these drugs is estimated to be 180 billion US dollars as of 2018<sup class="plainlinks noexcerpt noprint asof-tag update" style="display:none;"><a class="external text" href="https://en.wikipedia.org/w/index.php?title=G_protein-coupled_receptor&action=edit">[update]</a></sup>.<sup id="cite_ref-Hauser_2018_7-1" class="reference"><a href="#cite_note-Hauser_2018-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> It is estimated that GPCRs are targets for about 50% of drugs currently on the market, mainly due to their involvement in signaling pathways related to many diseases i.e. mental, metabolic including endocrinological disorders, immunological including viral infections, cardiovascular, inflammatory, senses disorders, and cancer. The long ago discovered association between GPCRs and many endogenous and exogenous substances, resulting in e.g. analgesia, is another dynamically developing field of the pharmaceutical research.<sup id="cite_ref-Trzaskowski2012_3-2" class="reference"><a href="#cite_note-Trzaskowski2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History_and_significance">History and significance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=1" title="Edit section: History and significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>With the determination of the first structure of the complex between a G-protein coupled receptor (GPCR) and a G-protein trimer (Gαβγ) in 2011 a new chapter of GPCR research was opened for structural investigations of global switches with more than one protein being investigated. The previous breakthroughs involved determination of the <a href="/wiki/Crystal_structure" title="Crystal structure">crystal structure</a> of the first GPCR, rhodopsin, in 2000 and the crystal structure of the first GPCR with a diffusible ligand (β<sub>2</sub>AR) in 2007. The way in which the seven transmembrane helices of a GPCR are arranged into a bundle was suspected based on the low-resolution model of frog rhodopsin from <a href="/wiki/Cryogenic_electron_microscopy" title="Cryogenic electron microscopy">cryogenic electron microscopy</a> studies of the two-dimensional crystals. The crystal structure of rhodopsin, that came up three years later, was not a surprise apart from the presence of an additional cytoplasmic helix H8 and a precise location of a loop covering retinal binding site. However, it provided a scaffold which was hoped to be a universal template for homology modeling and drug design for other GPCRs – a notion that proved to be too optimistic. </p><p>Seven years later, the crystallization of β<sub>2</sub>-adrenergic receptor (β<sub>2</sub>AR) with a diffusible ligand brought surprising results because it revealed quite a different shape of the receptor extracellular side than that of rhodopsin. This area is important because it is responsible for the ligand binding and is targeted by many drugs. Moreover, the ligand binding site was much more spacious than in the rhodopsin structure and was open to the exterior. In the other receptors crystallized shortly afterwards the binding side was even more easily accessible to the ligand. New structures complemented with biochemical investigations uncovered mechanisms of action of molecular switches which modulate the structure of the receptor leading to activation states for agonists or to complete or partial inactivation states for inverse agonists.<sup id="cite_ref-Trzaskowski2012_3-3" class="reference"><a href="#cite_note-Trzaskowski2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The 2012 <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a> was awarded to <a href="/wiki/Brian_Kobilka" title="Brian Kobilka">Brian Kobilka</a> and <a href="/wiki/Robert_Lefkowitz" title="Robert Lefkowitz">Robert Lefkowitz</a> for their work that was "crucial for understanding how G protein-coupled receptors function".<sup id="cite_ref-Nobel_committee,2012_8-0" class="reference"><a href="#cite_note-Nobel_committee,2012-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> There have been at least <a href="/wiki/List_of_Nobel_laureates_in_Physiology_or_Medicine" title="List of Nobel laureates in Physiology or Medicine">seven other Nobel Prizes</a> awarded for some aspect of G protein–mediated signaling. As of 2012, two of the top ten global best-selling drugs (<a href="/wiki/Fluticasone_propionate/salmeterol" class="mw-redirect" title="Fluticasone propionate/salmeterol">Advair Diskus</a> and <a href="/wiki/Aripiprazole" title="Aripiprazole">Abilify</a>) act by targeting G protein-coupled receptors.<sup id="cite_ref-urlwww.imshealth.com_9-0" class="reference"><a href="#cite_note-urlwww.imshealth.com-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=2" title="Edit section: Classification"><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:GPCR_classification.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/GPCR_classification.svg/220px-GPCR_classification.svg.png" decoding="async" width="220" height="97" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/GPCR_classification.svg/330px-GPCR_classification.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5b/GPCR_classification.svg/440px-GPCR_classification.svg.png 2x" data-file-width="384" data-file-height="169" /></a><figcaption>Classification Scheme of GPCRs in 2006. Since this time, more genes have been found. Class A (Rhodopsin-like), Class B (Secretin-like), Class C (Glutamate Receptor-like), Others (Adhesion (33), Frizzled (11), Taste type-2 (25), unclassified (23)).<sup id="cite_ref-pmid16753280_10-0" class="reference"><a href="#cite_note-pmid16753280-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The exact size of the GPCR superfamily is unknown, but at least 831 different <a href="/wiki/Human" title="Human">human</a> <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> (or about 4% of the entire <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein-coding</a> <a href="/wiki/Genome" title="Genome">genome</a>) have been predicted to code for them from genome <a href="/wiki/Sequence_analysis" title="Sequence analysis">sequence analysis</a>.<sup id="cite_ref-pmid16753280_10-1" class="reference"><a href="#cite_note-pmid16753280-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Although numerous classification schemes have been proposed, the superfamily was classically divided into three main classes (A, B, and C) with no detectable shared <a href="/wiki/Sequence_homology" title="Sequence homology">sequence homology</a> between classes. </p><p>The largest class by far is class A, which accounts for nearly 85% of the GPCR genes. Of class A GPCRs, over half of these are predicted to encode <a href="/wiki/Olfactory_receptor" title="Olfactory receptor">olfactory receptors</a>, while the remaining receptors are <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">liganded</a> by known <a href="/wiki/Endogenous" class="mw-redirect" title="Endogenous">endogenous</a> <a href="/wiki/Chemical_compound" title="Chemical compound">compounds</a> or are classified as <a href="/wiki/Orphan_receptor" title="Orphan receptor">orphan receptors</a>. Despite the lack of sequence homology between classes, all GPCRs have a common <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">structure</a> and mechanism of <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a>. The very large rhodopsin A group has been further subdivided into 19 subgroups (<a href="/wiki/Rhodopsin-like_receptors#Classes" title="Rhodopsin-like receptors">A1-A19</a>).<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>According to the classical A-F system, GPCRs can be grouped into six classes based on sequence homology and functional similarity:<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <ul><li><a href="/w/index.php?title=Class_A_GPCR&action=edit&redlink=1" class="new" title="Class A GPCR (page does not exist)">Class A</a> (or 1) (<a href="/wiki/Rhodopsin-like_receptors" title="Rhodopsin-like receptors">Rhodopsin-like</a>)</li> <li><a href="/wiki/Class_B_GPCR" class="mw-redirect" title="Class B GPCR">Class B</a> (or 2) (<a href="/wiki/Secretin_receptor_family" title="Secretin receptor family">Secretin receptor family</a>)</li> <li><a href="/wiki/Class_C_GPCR" title="Class C GPCR">Class C</a> (or 3) (<a href="/wiki/Metabotropic_glutamate_receptor" title="Metabotropic glutamate receptor">Metabotropic glutamate</a>/pheromone)</li> <li><a href="/w/index.php?title=Class_D_GPCR&action=edit&redlink=1" class="new" title="Class D GPCR (page does not exist)">Class D</a> (or 4) (<a href="/wiki/Fungal_mating_pheromone_receptors" title="Fungal mating pheromone receptors">Fungal mating pheromone receptors</a>)</li> <li><a href="/w/index.php?title=Class_E_GPCR&action=edit&redlink=1" class="new" title="Class E GPCR (page does not exist)">Class E</a> (or 5) (<a href="/wiki/Cyclic_AMP_receptors" title="Cyclic AMP receptors">Cyclic AMP receptors</a>)</li> <li><a href="/w/index.php?title=Class_F_GPCR&action=edit&redlink=1" class="new" title="Class F GPCR (page does not exist)">Class F</a> (or 6) (<a href="/wiki/Frizzled" title="Frizzled">Frizzled</a>/<a href="/wiki/Smoothened" title="Smoothened">Smoothened</a>)</li></ul> <p>More recently, an alternative classification system called <a href="/wiki/GRAFS" title="GRAFS">GRAFS</a> (<a href="/wiki/Metabotropic_glutamate_receptor" title="Metabotropic glutamate receptor">Glutamate</a>, <a href="/wiki/Rhodopsin" title="Rhodopsin">Rhodopsin</a>, <a href="/wiki/Adhesion_G_protein-coupled_receptor" title="Adhesion G protein-coupled receptor"><i>Adhesion</i></a>, <a href="/wiki/Frizzled" title="Frizzled">Frizzled</a>/<a href="/wiki/Taste_receptor" title="Taste receptor">Taste2</a>, <a href="/wiki/Secretin_receptor" title="Secretin receptor">Secretin</a>) has been proposed for vertebrate GPCRs.<sup id="cite_ref-pmid16753280_10-2" class="reference"><a href="#cite_note-pmid16753280-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> They correspond to classical classes C, A, B2, F, and B.<sup id="cite_ref-pmid22238661_17-0" class="reference"><a href="#cite_note-pmid22238661-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>An early study based on available DNA sequence suggested that the human genome encodes roughly 750 G protein-coupled receptors,<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> about 350 of which detect hormones, growth factors, and other endogenous ligands. Approximately 150 of the GPCRs found in the human genome have unknown functions. </p><p>Some web-servers<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> and bioinformatics prediction methods<sup id="cite_ref-pmid19364489_20-0" class="reference"><a href="#cite_note-pmid19364489-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19594431_21-0" class="reference"><a href="#cite_note-pmid19594431-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> have been used for predicting the classification of GPCRs according to their amino acid sequence alone, by means of the <a href="/wiki/Pseudo_amino_acid_composition" title="Pseudo amino acid composition">pseudo amino acid composition</a> approach. </p> <div class="mw-heading mw-heading2"><h2 id="Physiological_roles">Physiological roles</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=3" title="Edit section: Physiological roles"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>GPCRs are involved in a wide variety of physiological processes. Some examples of their physiological roles include: </p> <ol><li>The visual sense: The <a href="/wiki/Opsin" title="Opsin">opsins</a> use a <a href="/wiki/Photoisomerization" title="Photoisomerization">photoisomerization</a> reaction to translate <a href="/wiki/Electromagnetic_radiation" title="Electromagnetic radiation">electromagnetic radiation</a> into cellular signals. <a href="/wiki/Rhodopsin" title="Rhodopsin">Rhodopsin</a>, for example, uses the conversion of <a href="/wiki/Retinal" title="Retinal"><i>11-cis</i>-retinal</a> to <a href="/wiki/Retinal" title="Retinal"><i>all-trans</i>-retinal</a> for this purpose.</li> <li>The gustatory sense (taste): GPCRs in taste cells mediate release of <a href="/wiki/Gustducin" title="Gustducin">gustducin</a> in response to bitter-, umami- and sweet-tasting substances.</li> <li>The sense of smell: Receptors of the <a href="/wiki/Olfactory_epithelium" title="Olfactory epithelium">olfactory epithelium</a> bind odorants (olfactory receptors) and pheromones (vomeronasal receptors)</li> <li>Behavioral and mood regulation: Receptors in the <a href="/wiki/Mammal" title="Mammal">mammalian</a> <a href="/wiki/Brain" title="Brain">brain</a> bind several different <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitters</a>, including <a href="/wiki/Serotonin" title="Serotonin">serotonin</a>, <a href="/wiki/Dopamine" title="Dopamine">dopamine</a>, <a href="/wiki/Histamine" title="Histamine">histamine</a>, <a href="/wiki/Gamma-aminobutyric_acid" class="mw-redirect" title="Gamma-aminobutyric acid">GABA</a>, and <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a></li> <li>Regulation of <a href="/wiki/Immune_system" title="Immune system">immune system</a> activity and <a href="/wiki/Inflammation" title="Inflammation">inflammation</a>: <a href="/wiki/Chemokine" title="Chemokine">chemokine</a> receptors bind ligands that mediate intercellular communication between cells of the immune system; receptors such as <a href="/wiki/Histamine_receptor" title="Histamine receptor">histamine receptors</a> bind <a href="/wiki/Inflammatory_mediators" class="mw-redirect" title="Inflammatory mediators">inflammatory mediators</a> and engage target cell types in the <a href="/wiki/Inflammation" title="Inflammation">inflammatory response</a>. GPCRs are also involved in immune-modulation, e. g. regulating interleukin induction<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> or suppressing <a href="/wiki/Toll-like_receptor" title="Toll-like receptor">TLR</a>-induced immune responses from T cells.<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></li> <li>Autonomic nervous system transmission: Both the <a href="/wiki/Sympathetic_nervous_system" title="Sympathetic nervous system">sympathetic</a> and <a href="/wiki/Parasympathetic_nervous_system" title="Parasympathetic nervous system">parasympathetic</a> nervous systems are regulated by GPCR pathways, responsible for control of many automatic functions of the body such as blood pressure, heart rate, and digestive processes</li> <li>Cell density sensing: A novel GPCR role in regulating cell density sensing.</li> <li>Homeostasis modulation (e.g., water balance).<sup id="cite_ref-pmid21802439_24-0" class="reference"><a href="#cite_note-pmid21802439-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup></li> <li>Involved in growth and <a href="/wiki/Metastasis" title="Metastasis">metastasis</a> of some types of <a href="/wiki/Tumor" class="mw-redirect" title="Tumor">tumors</a>.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li> <li>Used in the endocrine system for peptide and amino-acid derivative hormones that bind to GCPRs on the cell membrane of a target cell. This activates cAMP, which in turn activates several kinases, allowing for a cellular response, such as transcription.</li></ol> <div class="mw-heading mw-heading2"><h2 id="Receptor_structure">Receptor structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=4" title="Edit section: Receptor structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>GPCRs are <a href="/wiki/Integral_membrane_protein" title="Integral membrane protein">integral membrane proteins</a> that possess seven membrane-spanning domains or <a href="/wiki/Transmembrane_helix" class="mw-redirect" title="Transmembrane helix">transmembrane helices</a>.<sup id="cite_ref-pmid_23407534_26-0" class="reference"><a href="#cite_note-pmid_23407534-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24359917_27-0" class="reference"><a href="#cite_note-pmid24359917-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The extracellular parts of the receptor can be <a href="/wiki/Glycosylation" title="Glycosylation">glycosylated</a>. These extracellular loops also contain two highly conserved <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> residues that form <a href="/wiki/Disulfide_bond" class="mw-redirect" title="Disulfide bond">disulfide bonds</a> to stabilize the receptor structure. Some seven-transmembrane helix proteins (<a href="/wiki/Channelrhodopsin" title="Channelrhodopsin">channelrhodopsin</a>) that resemble GPCRs may contain ion channels, within their protein. </p><p>In 2000, the first crystal structure of a mammalian GPCR, that of bovine <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a> (<span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1F88">1F88</a></span>), was solved.<sup id="cite_ref-pmid10926528_28-0" class="reference"><a href="#cite_note-pmid10926528-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In 2007, the first structure of a human GPCR was solved <sup id="cite_ref-Rasmussen_2007_29-0" class="reference"><a href="#cite_note-Rasmussen_2007-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cherezov_2007_1-1" class="reference"><a href="#cite_note-Cherezov_2007-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Rosenbaum_2007_30-0" class="reference"><a href="#cite_note-Rosenbaum_2007-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> This human <a href="/wiki/Beta-2_adrenergic_receptor" title="Beta-2 adrenergic receptor">β<sub>2</sub>-adrenergic receptor</a> GPCR structure proved highly similar to the bovine rhodopsin. The structures of activated or agonist-bound GPCRs have also been determined.<sup id="cite_ref-pmid21228869_31-0" class="reference"><a href="#cite_note-pmid21228869-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid21228876_32-0" class="reference"><a href="#cite_note-pmid21228876-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid21228877_33-0" class="reference"><a href="#cite_note-pmid21228877-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid21393508_34-0" class="reference"><a href="#cite_note-pmid21393508-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> These structures indicate how ligand binding at the extracellular side of a receptor leads to conformational changes in the cytoplasmic side of the receptor. The biggest change is an outward movement of the cytoplasmic part of the 5th and 6th transmembrane helix (TM5 and TM6). The structure of activated beta-2 adrenergic receptor in complex with G<sub>s</sub> confirmed that the Gα binds to a cavity created by this movement.<sup id="cite_ref-pmid21772288_35-0" class="reference"><a href="#cite_note-pmid21772288-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>GPCRs exhibit a similar structure to some other proteins with seven <a href="/wiki/Transmembrane_domain" title="Transmembrane domain">transmembrane domains</a>, such as <a href="/wiki/Microbial_rhodopsin" title="Microbial rhodopsin">microbial rhodopsins</a> and adiponectin receptors 1 and 2 (<a href="/wiki/ADIPOR1" class="mw-redirect" title="ADIPOR1">ADIPOR1</a> and <a href="/wiki/ADIPOR2" class="mw-redirect" title="ADIPOR2">ADIPOR2</a>). However, these 7TMH (7-transmembrane helices) receptors and channels do not associate with <a href="/wiki/G_protein" title="G protein">G proteins</a>. In addition, ADIPOR1 and ADIPOR2 are oriented oppositely to GPCRs in the membrane (i.e. GPCRs usually have an extracellular <a href="/wiki/N-terminus" title="N-terminus">N-terminus</a>, cytoplasmic <a href="/wiki/C-terminus" title="C-terminus">C-terminus</a>, whereas ADIPORs are inverted).<sup id="cite_ref-pmid12802337_36-0" class="reference"><a href="#cite_note-pmid12802337-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structure–function_relationships"><span id="Structure.E2.80.93function_relationships"></span>Structure–function relationships</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=5" title="Edit section: Structure–function relationships"><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:GPCR_in_membrane.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/GPCR_in_membrane.png/500px-GPCR_in_membrane.png" decoding="async" width="500" height="500" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/GPCR_in_membrane.png/750px-GPCR_in_membrane.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/GPCR_in_membrane.png/1000px-GPCR_in_membrane.png 2x" data-file-width="1667" data-file-height="1667" /></a><figcaption>Two-dimensional schematic of a generic GPCR set in a <a href="/wiki/Lipid_raft" title="Lipid raft">lipid raft</a>. Click the image for higher resolution to see details regarding the locations of important structures.</figcaption></figure> <p>In terms of structure, GPCRs are characterized by an extracellular <a href="/wiki/N-terminus" title="N-terminus">N-terminus</a>, followed by seven <a href="/wiki/Transmembrane_domain" title="Transmembrane domain">transmembrane</a> (7-TM) <a href="/wiki/Alpha_helix" title="Alpha helix">α-helices</a> (TM-1 to TM-7) connected by three intracellular (IL-1 to IL-3) and three extracellular loops (EL-1 to EL-3), and finally an intracellular <a href="/wiki/C-terminus" title="C-terminus">C-terminus</a>. The GPCR arranges itself into a <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">tertiary structure</a> resembling a barrel, with the seven transmembrane helices forming a cavity within the plasma membrane that serves a <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligand</a>-binding domain that is often covered by EL-2. Ligands may also bind elsewhere, however, as is the case for bulkier ligands (e.g., <a href="/wiki/Protein" title="Protein">proteins</a> or large <a href="/wiki/Peptide" title="Peptide">peptides</a>), which instead interact with the extracellular loops, or, as illustrated by the class C <a href="/wiki/Metabotropic_glutamate_receptors" class="mw-redirect" title="Metabotropic glutamate receptors">metabotropic glutamate receptors</a> (mGluRs), the N-terminal tail. The class C GPCRs are distinguished by their large N-terminal tail, which also contains a ligand-binding domain. Upon glutamate-binding to an mGluR, the N-terminal tail undergoes a conformational change that leads to its interaction with the residues of the extracellular loops and TM domains. The eventual effect of all three types of <a href="/wiki/Agonist" title="Agonist">agonist</a>-induced activation is a change in the relative orientations of the TM helices (likened to a twisting motion) leading to a wider intracellular surface and "revelation" of residues of the intracellular helices and TM domains crucial to signal transduction function (i.e., G-protein coupling). <a href="/wiki/Inverse_agonist" title="Inverse agonist">Inverse agonists</a> and <a href="/wiki/Receptor_antagonist" title="Receptor antagonist">antagonists</a> may also bind to a number of different sites, but the eventual effect must be prevention of this TM helix reorientation.<sup id="cite_ref-Trzaskowski2012_3-4" class="reference"><a href="#cite_note-Trzaskowski2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The structure of the N- and C-terminal tails of GPCRs may also serve important functions beyond ligand-binding. For example, The C-terminus of M<sub>3</sub> muscarinic receptors is sufficient, and the six-amino-acid polybasic (KKKRRK) domain in the C-terminus is necessary for its preassembly with G<sub>q</sub> proteins.<sup id="cite_ref-pmid=21873996_37-0" class="reference"><a href="#cite_note-pmid=21873996-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> In particular, the C-terminus often contains <a href="/wiki/Serine" title="Serine">serine</a> (Ser) or <a href="/wiki/Threonine" title="Threonine">threonine</a> (Thr) residues that, when <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylated</a>, increase the <a href="/wiki/Affinity_(pharmacology)" class="mw-redirect" title="Affinity (pharmacology)">affinity</a> of the intracellular surface for the binding of scaffolding proteins called β-<a href="/wiki/Arrestin" title="Arrestin">arrestins</a> (β-arr).<sup id="cite_ref-pmid2163110_38-0" class="reference"><a href="#cite_note-pmid2163110-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Once bound, β-arrestins both <a href="/wiki/Sterically" class="mw-redirect" title="Sterically">sterically</a> prevent G-protein coupling and may recruit other proteins, leading to the creation of signaling complexes involved in extracellular-signal regulated kinase (<a href="/wiki/Extracellular_signal-regulated_kinases" title="Extracellular signal-regulated kinases">ERK</a>) pathway activation or receptor <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a> (internalization). As the phosphorylation of these Ser and Thr residues often occurs as a result of GPCR activation, the β-arr-mediated G-protein-decoupling and internalization of GPCRs are important mechanisms of <a href="/wiki/Desensitization_(medicine)" title="Desensitization (medicine)">desensitization</a>.<sup id="cite_ref-pmid11861753_39-0" class="reference"><a href="#cite_note-pmid11861753-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> In addition, internalized "mega-complexes" consisting of a single GPCR, β-arr(in the tail conformation),<sup id="cite_ref-pmid28223524_40-0" class="reference"><a href="#cite_note-pmid28223524-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid27827372_41-0" class="reference"><a href="#cite_note-pmid27827372-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> and heterotrimeric G protein exist and may account for protein signaling from endosomes.<sup id="cite_ref-pmid27499021_42-0" class="reference"><a href="#cite_note-pmid27499021-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid31740855_43-0" class="reference"><a href="#cite_note-pmid31740855-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>A final common structural theme among GPCRs is <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> of one or more sites of the C-terminal tail or the intracellular loops. Palmitoylation is the covalent modification of <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> (Cys) residues via addition of hydrophobic <a href="/wiki/Acyl_group" title="Acyl group">acyl groups</a>, and has the effect of targeting the receptor to <a href="/wiki/Cholesterol" title="Cholesterol">cholesterol</a>- and <a href="/wiki/Sphingolipid" title="Sphingolipid">sphingolipid</a>-rich microdomains of the plasma membrane called <a href="/wiki/Lipid_raft" title="Lipid raft">lipid rafts</a>. As many of the downstream transducer and effector molecules of GPCRs (including those involved in <a href="/wiki/Negative_feedback" title="Negative feedback">negative feedback</a> pathways) are also targeted to lipid rafts, this has the effect of facilitating rapid receptor signaling. </p><p>GPCRs respond to extracellular signals mediated by a huge diversity of agonists, ranging from proteins to <a href="/wiki/Biogenic_amines" class="mw-redirect" title="Biogenic amines">biogenic amines</a> to <a href="/wiki/Protons" class="mw-redirect" title="Protons">protons</a>, but all transduce this signal via a mechanism of G-protein coupling. This is made possible by a <a href="/wiki/Guanine" title="Guanine">guanine</a>-nucleotide exchange factor (<a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">GEF</a>) domain primarily formed by a combination of IL-2 and IL-3 along with adjacent residues of the associated TM helices. </p> <div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=6" title="Edit section: Mechanism"><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:GPCR_activation.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/GPCR_activation.jpg/400px-GPCR_activation.jpg" decoding="async" width="400" height="400" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/GPCR_activation.jpg/600px-GPCR_activation.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e6/GPCR_activation.jpg/800px-GPCR_activation.jpg 2x" data-file-width="1667" data-file-height="1667" /></a><figcaption>Cartoon depicting the basic concept of GPCR conformational activation. Ligand binding disrupts an ionic lock between the E/DRY motif of TM-3 and acidic residues of TM-6. As a result, the GPCR reorganizes to allow activation of G-alpha proteins. The "side perspective" is a view from above and to the side of the GPCR as it is set in the plasma membrane (the membrane lipids have been omitted for clarity). The incorrectly labelled "intracellular perspective" shows an extracellular view looking down at the plasma membrane from outside the cell.<sup id="cite_ref-pmid20019124_44-0" class="reference"><a href="#cite_note-pmid20019124-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The G protein-coupled receptor is activated by an external signal in the form of a ligand or other signal mediator. This creates a conformational change in the receptor, causing activation of a <a href="/wiki/G_protein" title="G protein">G protein</a>. Further effect depends on the type of G protein. G proteins are subsequently inactivated by GTPase activating proteins, known as <a href="/wiki/Regulator_of_G_protein_signaling" title="Regulator of G protein signaling">RGS proteins</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Ligand_binding">Ligand binding</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=7" title="Edit section: Ligand binding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>GPCRs include one or more receptors for the following ligands: sensory signal mediators (e.g., light and <a href="/wiki/Olfactory" class="mw-redirect" title="Olfactory">olfactory</a> stimulatory molecules); <a href="/wiki/Adenosine" title="Adenosine">adenosine</a>, <a href="/wiki/Bombesin" title="Bombesin">bombesin</a>, <a href="/wiki/Bradykinin" title="Bradykinin">bradykinin</a>, <a href="/wiki/Endothelin" title="Endothelin">endothelin</a>, γ-aminobutyric acid (<a href="/wiki/Gamma-aminobutyric_acid" class="mw-redirect" title="Gamma-aminobutyric acid">GABA</a>), hepatocyte growth factor (<a href="/wiki/Hepatocyte_growth_factor" title="Hepatocyte growth factor">HGF</a>), <a href="/wiki/Melanocortin" title="Melanocortin">melanocortins</a>, <a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">neuropeptide Y</a>, <a href="/wiki/Opioid" title="Opioid">opioid</a> peptides, <a href="/wiki/Opsin" title="Opsin">opsins</a>, <a href="/wiki/Somatostatin" title="Somatostatin">somatostatin</a>, <a href="/wiki/Growth_hormone" title="Growth hormone">GH</a>, <a href="/wiki/Tachykinins" class="mw-redirect" title="Tachykinins">tachykinins</a>, members of the <a href="/wiki/Vasoactive_intestinal_peptide" title="Vasoactive intestinal peptide">vasoactive intestinal peptide</a> family, and <a href="/wiki/Vasopressin" title="Vasopressin">vasopressin</a>; <a href="/wiki/Biogenic_amine" title="Biogenic amine">biogenic amines</a> (e.g., <a href="/wiki/Dopamine" title="Dopamine">dopamine</a>, <a href="/wiki/Epinephrine" class="mw-redirect" title="Epinephrine">epinephrine</a>, <a href="/wiki/Norepinephrine" title="Norepinephrine">norepinephrine</a>, <a href="/wiki/Histamine" title="Histamine">histamine</a>, <a href="/wiki/Serotonin" title="Serotonin">serotonin</a>, and <a href="/wiki/Melatonin" title="Melatonin">melatonin</a>); <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> (<a href="/wiki/Metabotropic" class="mw-redirect" title="Metabotropic">metabotropic</a> effect); <a href="/wiki/Glucagon" title="Glucagon">glucagon</a>; <a href="/wiki/Acetylcholine" title="Acetylcholine">acetylcholine</a> (<a href="/wiki/Muscarinic" class="mw-redirect" title="Muscarinic">muscarinic</a> effect); <a href="/wiki/Chemokines" class="mw-redirect" title="Chemokines">chemokines</a>; <a href="/wiki/Lipid" title="Lipid">lipid</a> mediators of <a href="/wiki/Inflammation" title="Inflammation">inflammation</a> (e.g., <a href="/wiki/Prostaglandins" class="mw-redirect" title="Prostaglandins">prostaglandins</a>, <a href="/wiki/Prostanoid" title="Prostanoid">prostanoids</a>, <a href="/wiki/Platelet-activating_factor" title="Platelet-activating factor">platelet-activating factor</a>, and <a href="/wiki/Leukotrienes" class="mw-redirect" title="Leukotrienes">leukotrienes</a>); peptide hormones (e.g., <a href="/wiki/Calcitonin" title="Calcitonin">calcitonin</a>, C5a <a href="/wiki/Anaphylatoxin" title="Anaphylatoxin">anaphylatoxin</a>, <a href="/wiki/Follicle-stimulating_hormone" title="Follicle-stimulating hormone">follicle-stimulating hormone</a> [FSH], <a href="/wiki/Gonadotropin-releasing_hormone" title="Gonadotropin-releasing hormone">gonadotropin-releasing hormone</a> [GnRH], <a href="/wiki/Neurokinin" class="mw-redirect" title="Neurokinin">neurokinin</a>, <a href="/wiki/Thyrotropin-releasing_hormone" title="Thyrotropin-releasing hormone">thyrotropin-releasing hormone</a> [TRH], and <a href="/wiki/Oxytocin" title="Oxytocin">oxytocin</a>); and <a href="/wiki/Endocannabinoid" class="mw-redirect" title="Endocannabinoid">endocannabinoids</a>. </p><p>GPCRs that act as receptors for stimuli that have not yet been identified are known as <a href="/wiki/Orphan_receptor" title="Orphan receptor">orphan receptors</a>. </p><p>However, in contrast to other types of receptors that have been studied, wherein ligands bind externally to the membrane, the <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligands</a> of GPCRs typically bind within the transmembrane domain. However, <a href="/wiki/Protease-activated_receptor" title="Protease-activated receptor">protease-activated receptors</a> are activated by cleavage of part of their extracellular domain.<sup id="cite_ref-pmid12970120_45-0" class="reference"><a href="#cite_note-pmid12970120-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Conformational_change">Conformational change</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=8" title="Edit section: Conformational change"><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:Beta2Receptor-with-Gs.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Beta2Receptor-with-Gs.png/300px-Beta2Receptor-with-Gs.png" decoding="async" width="300" height="366" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Beta2Receptor-with-Gs.png/450px-Beta2Receptor-with-Gs.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/Beta2Receptor-with-Gs.png/600px-Beta2Receptor-with-Gs.png 2x" data-file-width="806" data-file-height="984" /></a><figcaption>Crystal structure of activated beta-2 adrenergic receptor in complex with G<sub>s</sub>(<a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> entry <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180128134132/https://www.rcsb.org/structure/3SN6">3SN6</a>). The receptor is colored red, Gα green, Gβ cyan, and Gγ yellow. The C-terminus of Gα is located in a cavity created by an outward movement of the cytoplasmic parts of TM5 and 6.</figcaption></figure> <p>The <a href="/wiki/Signal_transduction" title="Signal transduction">transduction of the signal</a> through the membrane by the receptor is not completely understood. It is known that in the inactive state, the GPCR is bound to a <a href="/wiki/Heterotrimeric_G_protein" title="Heterotrimeric G protein">heterotrimeric G protein</a> complex. Binding of an agonist to the GPCR results in a <a href="/wiki/Conformational_change" title="Conformational change">conformational change</a> in the receptor that is transmitted to the bound G<sub>α</sub> subunit of the heterotrimeric G protein via <a href="/wiki/Protein_dynamics#Global_flexibility:_multiple_domains" title="Protein dynamics">protein domain dynamics</a>. The activated G<sub>α</sub> subunit exchanges <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> in place of <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">GDP</a> which in turn triggers the dissociation of G<sub>α</sub> subunit from the G<sub>βγ</sub> dimer and from the receptor. The dissociated G<sub>α</sub> and G<sub>βγ</sub> subunits interact with other intracellular proteins to continue the signal transduction cascade while the freed GPCR is able to rebind to another heterotrimeric G protein to form a new complex that is ready to initiate another round of signal transduction.<sup id="cite_ref-pmid17095603_46-0" class="reference"><a href="#cite_note-pmid17095603-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>It is believed that a receptor molecule exists in a conformational <a href="/wiki/Dynamic_equilibrium" title="Dynamic equilibrium">equilibrium</a> between active and inactive biophysical states.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> The binding of ligands to the receptor may shift the equilibrium toward the active receptor states. Three types of ligands exist: Agonists are ligands that shift the equilibrium in favour of active states; <a href="/wiki/Inverse_agonist" title="Inverse agonist">inverse agonists</a> are ligands that shift the equilibrium in favour of inactive states; and neutral antagonists are ligands that do not affect the equilibrium. It is not yet known how exactly the active and inactive states differ from each other. </p> <div class="mw-heading mw-heading3"><h3 id="G-protein_activation/deactivation_cycle"><span id="G-protein_activation.2Fdeactivation_cycle"></span>G-protein activation/deactivation cycle</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=9" title="Edit section: G-protein activation/deactivation cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:GPCR_cycle.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/GPCR_cycle.jpg/500px-GPCR_cycle.jpg" decoding="async" width="500" height="375" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/GPCR_cycle.jpg/750px-GPCR_cycle.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c9/GPCR_cycle.jpg/1000px-GPCR_cycle.jpg 2x" data-file-width="1667" data-file-height="1250" /></a><figcaption>Cartoon depicting the heterotrimeric G-protein activation/deactivation cycle in the context of GPCR signaling</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/G_protein" title="G protein">G protein</a></div> <p>When the receptor is inactive, the <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">GEF</a> domain may be bound to an also inactive α-subunit of a <a href="/wiki/Heterotrimeric_G-protein" class="mw-redirect" title="Heterotrimeric G-protein">heterotrimeric G-protein</a>. These "G-proteins" are a <a href="/wiki/Protein_trimer" title="Protein trimer">trimer</a> of α, β, and γ subunits (known as Gα, Gβ, and Gγ, respectively) that is rendered inactive when reversibly bound to <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">Guanosine diphosphate</a> (GDP) (or, alternatively, no guanine nucleotide) but active when bound to <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">guanosine triphosphate</a> (GTP). Upon receptor activation, the GEF domain, in turn, <a href="/wiki/Allosterically" class="mw-redirect" title="Allosterically">allosterically</a> activates the G-protein by facilitating the exchange of a molecule of GDP for GTP at the G-protein's α-subunit. The cell maintains a 10:1 ratio of cytosolic GTP:GDP so exchange for GTP is ensured. At this point, the subunits of the G-protein dissociate from the receptor, as well as each other, to yield a Gα-GTP <a href="/wiki/Monomer" title="Monomer">monomer</a> and a tightly interacting <a href="/wiki/G_beta-gamma_complex" title="G beta-gamma complex">Gβγ dimer</a>, which are now free to modulate the activity of other intracellular proteins. The extent to which they may <a href="/wiki/Diffuse" class="mw-redirect" title="Diffuse">diffuse</a>, however, is limited due to the <a href="/wiki/Palmitoylation" title="Palmitoylation">palmitoylation</a> of Gα and the presence of an <a href="/wiki/Isoprenoid" class="mw-redirect" title="Isoprenoid">isoprenoid</a> moiety that has been <a href="/wiki/Covalent_bond" title="Covalent bond">covalently</a> added to the C-termini of Gγ. </p><p>Because Gα also has slow <a href="/wiki/GTP-ase" class="mw-redirect" title="GTP-ase">GTP→GDP hydrolysis</a> capability, the inactive form of the α-subunit (Gα-GDP) is eventually regenerated, thus allowing reassociation with a Gβγ dimer to form the "resting" G-protein, which can again bind to a GPCR and await activation. The rate of GTP hydrolysis is often accelerated due to the actions of another family of allosteric modulating proteins called <a href="/wiki/Regulator_of_G_protein_signaling" title="Regulator of G protein signaling">regulators of G-protein signaling</a>, or RGS proteins, which are a type of <a href="/wiki/GTPase-activating_protein" title="GTPase-activating protein">GTPase-activating protein</a>, or GAP. In fact, many of the primary <a href="/wiki/Effector_(biology)" title="Effector (biology)">effector</a> proteins (e.g., <a href="/wiki/Adenylate_cyclase" class="mw-redirect" title="Adenylate cyclase">adenylate cyclases</a>) that become activated/inactivated upon interaction with Gα-GTP also have GAP activity. Thus, even at this early stage in the process, GPCR-initiated signaling has the capacity for self-termination. </p> <div class="mw-heading mw-heading3"><h3 id="Crosstalk">Crosstalk</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=10" title="Edit section: Crosstalk"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:GPCR_and_itegrin_signaling_diagram.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/GPCR_and_itegrin_signaling_diagram.png/220px-GPCR_and_itegrin_signaling_diagram.png" decoding="async" width="220" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/GPCR_and_itegrin_signaling_diagram.png/330px-GPCR_and_itegrin_signaling_diagram.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/34/GPCR_and_itegrin_signaling_diagram.png/440px-GPCR_and_itegrin_signaling_diagram.png 2x" data-file-width="819" data-file-height="846" /></a><figcaption>Proposed downstream interactions between <a href="/wiki/Integrin" title="Integrin">integrin</a> signaling and GPCRs. Integrins are shown elevating Ca<sup>2+</sup> and phosphorylating FAK, which is weakening GPCR signaling.</figcaption></figure> <p>GPCRs downstream signals have been shown to possibly interact with <a href="/wiki/Integrin" title="Integrin">integrin</a> signals, such as <a href="/wiki/PTK2" title="PTK2">FAK</a>.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Integrin signaling will phosphorylate FAK, which can then decrease GPCR G<sub>αs</sub> activity. </p> <div class="mw-heading mw-heading2"><h2 id="Signaling">Signaling</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=11" title="Edit section: Signaling"><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:GPCR_mechanism.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/GPCR_mechanism.png/300px-GPCR_mechanism.png" decoding="async" width="300" height="612" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/GPCR_mechanism.png/450px-GPCR_mechanism.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/90/GPCR_mechanism.png/600px-GPCR_mechanism.png 2x" data-file-width="766" data-file-height="1563" /></a><figcaption>G-protein-coupled receptor mechanism</figcaption></figure> <p>If a receptor in an active state encounters a <a href="/wiki/G_protein" title="G protein">G protein</a>, it may activate it. Some evidence suggests that receptors and G proteins are actually pre-coupled.<sup id="cite_ref-pmid=21873996_37-1" class="reference"><a href="#cite_note-pmid=21873996-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> For example, binding of G proteins to receptors affects the receptor's affinity for ligands. Activated G proteins are bound to <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a>. </p><p>Further signal transduction depends on the type of G protein. The enzyme <a href="/wiki/Adenylate_cyclase" class="mw-redirect" title="Adenylate cyclase">adenylate cyclase</a> is an example of a cellular protein that can be regulated by a G protein, in this case the G protein <a href="/wiki/Gs_alpha_subunit" title="Gs alpha subunit">G<sub>s</sub></a>. Adenylate cyclase activity is activated when it binds to a subunit of the activated G protein. Activation of adenylate cyclase ends when the G protein returns to the <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">GDP</a>-bound state. </p><p>Adenylate cyclases (of which 9 membrane-bound and one cytosolic forms are known in humans) may also be activated or inhibited in other ways (e.g., Ca2+/<a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a> binding), which can modify the activity of these enzymes in an additive or synergistic fashion along with the G proteins. </p><p>The signaling pathways activated through a GPCR are limited by the <a href="/wiki/Protein_primary_structure" title="Protein primary structure">primary sequence</a> and <a href="/wiki/Tertiary_structure" class="mw-redirect" title="Tertiary structure">tertiary structure</a> of the GPCR itself but ultimately determined by the particular <a href="/wiki/Protein_conformation" class="mw-redirect" title="Protein conformation">conformation</a> stabilized by a particular <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligand</a>, as well as the availability of <a href="/wiki/Transducer" title="Transducer">transducer</a> molecules. Currently, GPCRs are considered to utilize two primary types of transducers: <a href="/wiki/G-proteins" class="mw-redirect" title="G-proteins">G-proteins</a> and <a href="/wiki/Arrestin" title="Arrestin">β-arrestins</a>. Because β-arr's have high <a href="/wiki/Affinity_(pharmacology)" class="mw-redirect" title="Affinity (pharmacology)">affinity</a> only to the <a href="/wiki/Phosphorylated" class="mw-redirect" title="Phosphorylated">phosphorylated</a> form of most GPCRs (see above or below), the majority of signaling is ultimately dependent upon G-protein activation. However, the possibility for interaction does allow for G-protein-independent signaling to occur. </p> <div class="mw-heading mw-heading3"><h3 id="G-protein-dependent_signaling">G-protein-dependent signaling</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=12" title="Edit section: G-protein-dependent signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are three main G-protein-mediated signaling pathways, mediated by four <a href="/wiki/Class_(biology)" title="Class (biology)">sub-classes</a> of G-proteins distinguished from each other by <a href="/wiki/Sequence_homology" title="Sequence homology">sequence homology</a> (<a href="/wiki/G%CE%B1s" class="mw-redirect" title="Gαs">G<sub>αs</sub></a>, <a href="/wiki/G%CE%B1i" class="mw-redirect" title="Gαi">G<sub>αi/o</sub></a>, <a href="/wiki/G%CE%B1q" class="mw-redirect" title="Gαq">G<sub>αq/11</sub></a>, and <a href="/wiki/G12/G13_alpha_subunits" title="G12/G13 alpha subunits">G<sub>α12/13</sub></a>). Each sub-class of G-protein consists of multiple proteins, each the product of multiple <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> or <a href="/wiki/Splice_variant" class="mw-redirect" title="Splice variant">splice variations</a> that may imbue them with differences ranging from subtle to distinct with regard to signaling properties, but in general they appear reasonably grouped into four classes. Because the signal transducing properties of the various possible <a href="/wiki/G_beta-gamma_complex" title="G beta-gamma complex">βγ combinations</a> do not appear to radically differ from one another, these classes are defined according to the isoform of their α-subunit.<sup id="cite_ref-Wettschureck_2005_6-1" class="reference"><a href="#cite_note-Wettschureck_2005-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 1163">: 1163 </span></sup> </p><p>While most GPCRs are capable of activating more than one Gα-subtype, they also show a preference for one subtype over another. When the subtype activated depends on the ligand that is bound to the GPCR, this is called <a href="/wiki/Functional_selectivity" title="Functional selectivity">functional selectivity</a> (also known as agonist-directed trafficking, or conformation-specific agonism). However, the binding of any single particular <a href="/wiki/Agonist" title="Agonist">agonist</a> may also initiate activation of multiple different G-proteins, as it may be capable of stabilizing more than one conformation of the GPCR's <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">GEF</a> domain, even over the course of a single interaction. In addition, a conformation that preferably activates one <a href="/wiki/Isoform" class="mw-redirect" title="Isoform">isoform</a> of Gα may activate another if the preferred is less available. Furthermore, <a href="/wiki/Feedback" title="Feedback">feedback</a> pathways may result in <a href="/wiki/Post-translational_modification" title="Post-translational modification">receptor modifications</a> (e.g., phosphorylation) that alter the G-protein preference. Regardless of these various nuances, the GPCR's preferred coupling partner is usually defined according to the G-protein most obviously activated by the <a href="/wiki/Endogenous" class="mw-redirect" title="Endogenous">endogenous</a> ligand under most <a href="/wiki/Physiological" class="mw-redirect" title="Physiological">physiological</a> or <a href="/wiki/Experimental" class="mw-redirect" title="Experimental">experimental</a> conditions. </p> <div class="mw-heading mw-heading4"><h4 id="Gα_signaling"><span id="G.CE.B1_signaling"></span>Gα signaling</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=13" title="Edit section: Gα signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ol><li>The effector of both the G<sub>αs</sub> and G<sub>αi/o</sub> pathways is the <a href="/wiki/Cyclic_amp" class="mw-redirect" title="Cyclic amp">cyclic-adenosine monophosphate</a> (cAMP)-generating enzyme <a href="/wiki/Adenylyl_cyclase" title="Adenylyl cyclase">adenylate cyclase</a>, or AC. While there are ten different AC gene products in mammals, each with subtle differences in <a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissue</a> distribution or function, all <a href="/wiki/Catalyze" class="mw-redirect" title="Catalyze">catalyze</a> the conversion of <a href="/wiki/Cytosolic" class="mw-redirect" title="Cytosolic">cytosolic</a> <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP) to cAMP, and all are directly stimulated by G-proteins of the G<sub>αs</sub> class. In contrast, however, interaction with Gα subunits of the G<sub>αi/o</sub> type inhibits AC from generating cAMP. Thus, a GPCR coupled to G<sub>αs</sub> counteracts the actions of a GPCR coupled to G<sub>αi/o</sub>, and vice versa. The level of cytosolic cAMP may then determine the activity of various <a href="/wiki/Cyclic_nucleotide-gated_ion_channel" class="mw-redirect" title="Cyclic nucleotide-gated ion channel">ion channels</a> as well as members of the <a href="/wiki/Serine/threonine-specific_protein_kinase" title="Serine/threonine-specific protein kinase">ser/thr-specific</a> <a href="/wiki/Protein_kinase_A" title="Protein kinase A">protein kinase A</a> (PKA) family. Thus cAMP is considered a <a href="/wiki/Second_messenger_system" title="Second messenger system">second messenger</a> and PKA a secondary <a href="/wiki/Effector_(biology)" title="Effector (biology)">effector</a>.</li> <li>The effector of the G<sub>αq/11</sub> pathway is <a href="/wiki/Phospholipase_C" title="Phospholipase C">phospholipase C-β</a> (PLCβ), which catalyzes the cleavage of membrane-bound <a href="/wiki/Phosphatidylinositol_4,5-bisphosphate" title="Phosphatidylinositol 4,5-bisphosphate">phosphatidylinositol 4,5-bisphosphate</a> (PIP2) into the second messengers <a href="/wiki/Inositol_trisphosphate" title="Inositol trisphosphate">inositol (1,4,5) trisphosphate</a> (IP3) and <a href="/wiki/Diglyceride" title="Diglyceride">diacylglycerol</a> (DAG). IP3 acts on <a href="/wiki/Inositol_trisphosphate_receptor" title="Inositol trisphosphate receptor">IP3 receptors</a> found in the membrane of the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a> (ER) to elicit <a href="/wiki/Ca2%2B" class="mw-redirect" title="Ca2+">Ca<sup>2+</sup></a> release from the ER, while DAG diffuses along the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> where it may activate any membrane localized forms of a second ser/thr kinase called <a href="/wiki/Protein_kinase_C" title="Protein kinase C">protein kinase C</a> (PKC). Since many isoforms of PKC are also activated by increases in intracellular Ca<sup>2+</sup>, both these pathways can also converge on each other to signal through the same secondary effector. Elevated intracellular Ca<sup>2+</sup> also binds and <a href="/wiki/Allosterically" class="mw-redirect" title="Allosterically">allosterically</a> activates proteins called <a href="/wiki/Calmodulin" title="Calmodulin">calmodulins</a>, which in turn tosolic <a href="/wiki/Small_GTPase" title="Small GTPase">small GTPase</a>, <a href="/wiki/Rho_family_of_GTPases" title="Rho family of GTPases">Rho</a>. Once bound to GTP, Rho can then go on to activate various proteins responsible for <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a> regulation such as <a href="/wiki/Rho-associated_protein_kinase" title="Rho-associated protein kinase">Rho-kinase</a> (ROCK). Most GPCRs that couple to G<sub>α12/13</sub> also couple to other sub-classes, often G<sub>αq/11</sub>.</li></ol> <div class="mw-heading mw-heading4"><h4 id="Gβγ_signaling"><span id="G.CE.B2.CE.B3_signaling"></span>Gβγ signaling</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=14" title="Edit section: Gβγ signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The above descriptions ignore the effects of <a href="/wiki/G_beta-gamma_complex" title="G beta-gamma complex">Gβγ</a>–signalling, which can also be important, in particular in the case of activated G<sub>αi/o</sub>-coupled GPCRs. The primary effectors of Gβγ are various ion channels, such as <a href="/wiki/G_protein-coupled_inwardly-rectifying_potassium_channel" class="mw-redirect" title="G protein-coupled inwardly-rectifying potassium channel">G-protein-regulated inwardly rectifying K<sup>+</sup> channels</a> (GIRKs), <a href="/wiki/P-type_calcium_channel" title="P-type calcium channel">P</a>/<a href="/wiki/Q-type_calcium_channel" title="Q-type calcium channel">Q</a>- and <a href="/wiki/N-type_calcium_channel" title="N-type calcium channel">N-type</a> <a href="/wiki/Voltage-dependent_calcium_channel" class="mw-redirect" title="Voltage-dependent calcium channel">voltage-gated Ca<sup>2+</sup> channels</a>, as well as some isoforms of AC and PLC, along with some <a href="/wiki/PI3K" class="mw-redirect" title="PI3K">phosphoinositide-3-kinase</a> (PI3K) isoforms. </p> <div class="mw-heading mw-heading3"><h3 id="G-protein-independent_signaling">G-protein-independent signaling</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=15" title="Edit section: G-protein-independent signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although they are classically thought of working only together, GPCRs may signal through G-protein-independent mechanisms, and heterotrimeric G-proteins may play functional roles independent of GPCRs. GPCRs may signal independently through many proteins already mentioned for their roles in G-protein-dependent signaling such as <a href="/wiki/Arrestin" title="Arrestin">β-arrs</a>, <a href="/wiki/G_protein-coupled_receptor_kinase" title="G protein-coupled receptor kinase">GRKs</a>, and <a href="/wiki/Src_(gene)" class="mw-redirect" title="Src (gene)">Srcs</a>. Such signaling has been shown to be physiologically relevant, for example, <a href="/wiki/Arrestin" title="Arrestin">β-arrestin</a> signaling mediated by the chemokine receptor <a href="/wiki/CXCR3" title="CXCR3">CXCR3</a> was necessary for full efficacy chemotaxis of activated T cells.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> In addition, further scaffolding proteins involved in <a href="/wiki/Subcellular_localization" title="Subcellular localization">subcellular localization</a> of GPCRs (e.g., <a href="/wiki/PDZ_(biology)" class="mw-redirect" title="PDZ (biology)">PDZ-domain</a>-containing proteins) may also act as signal transducers. Most often the effector is a member of the <a href="/wiki/MAPK" class="mw-redirect" title="MAPK">MAPK</a> family. </p> <div class="mw-heading mw-heading4"><h4 id="Examples">Examples</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=16" title="Edit section: Examples"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the late 1990s, evidence began accumulating to suggest that some GPCRs are able to signal without G proteins. The <a href="/wiki/MAPK1" title="MAPK1">ERK2</a> mitogen-activated protein kinase, a key signal transduction mediator downstream of receptor activation in many pathways, has been shown to be activated in response to cAMP-mediated receptor activation in the <a href="/wiki/Slime_mold" title="Slime mold">slime mold</a> <a href="/wiki/Dictyostelium_discoideum" title="Dictyostelium discoideum"><i>D. discoideum</i></a> despite the absence of the associated G protein α- and β-subunits.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>In mammalian cells, the much-studied β<sub>2</sub>-adrenoceptor has been demonstrated to activate the ERK2 pathway after arrestin-mediated uncoupling of G-protein-mediated signaling. Therefore, it seems likely that some mechanisms previously believed related purely to receptor desensitisation are actually examples of receptors switching their signaling pathway, rather than simply being switched off. </p><p>In kidney cells, the <a href="/wiki/Bradykinin_receptor_B2" title="Bradykinin receptor B2">bradykinin receptor B2</a> has been shown to interact directly with a protein tyrosine phosphatase. The presence of a tyrosine-phosphorylated <a href="/wiki/Immunoreceptor_tyrosine-based_inhibitory_motif" title="Immunoreceptor tyrosine-based inhibitory motif">ITIM</a> (immunoreceptor tyrosine-based inhibitory motif) sequence in the B2 receptor is necessary to mediate this interaction and subsequently the antiproliferative effect of bradykinin.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="GPCR-independent_signaling_by_heterotrimeric_G-proteins">GPCR-independent signaling by heterotrimeric G-proteins</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=17" title="Edit section: GPCR-independent signaling by heterotrimeric G-proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although it is a relatively immature area of research, it appears that heterotrimeric G-proteins may also take part in non-GPCR signaling. There is evidence for roles as signal transducers in nearly all other types of receptor-mediated signaling, including <a href="/wiki/Integrins" class="mw-redirect" title="Integrins">integrins</a>, <a href="/wiki/Receptor_tyrosine_kinases" class="mw-redirect" title="Receptor tyrosine kinases">receptor tyrosine kinases</a> (RTKs), <a href="/wiki/Cytokine_receptors" class="mw-redirect" title="Cytokine receptors">cytokine receptors</a> (<a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK/STATs</a>), as well as modulation of various other "accessory" proteins such as <a href="/wiki/Guanine_nucleotide_exchange_factor" title="Guanine nucleotide exchange factor">GEFs</a>, <a href="/wiki/Guanosine_nucleotide_dissociation_inhibitors" class="mw-redirect" title="Guanosine nucleotide dissociation inhibitors">guanine-nucleotide dissociation inhibitors</a> (GDIs) and <a href="/wiki/Protein_phosphatases" class="mw-redirect" title="Protein phosphatases">protein phosphatases</a>. There may even be specific proteins of these classes whose primary function is as part of GPCR-independent pathways, termed activators of G-protein signalling (AGS). Both the ubiquity of these interactions and the importance of Gα vs. Gβγ subunits to these processes are still unclear. </p> <div class="mw-heading mw-heading2"><h2 id="Details_of_cAMP_and_PIP2_pathways">Details of cAMP and PIP2 pathways</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=18" title="Edit section: Details of cAMP and PIP2 pathways"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Proteinkinase_1.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Proteinkinase_1.svg/250px-Proteinkinase_1.svg.png" decoding="async" width="250" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Proteinkinase_1.svg/375px-Proteinkinase_1.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/66/Proteinkinase_1.svg/500px-Proteinkinase_1.svg.png 2x" data-file-width="543" data-file-height="305" /></a><figcaption>Activation effects of cAMP on protein kinase A</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg/250px-The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg" decoding="async" width="250" height="174" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg/375px-The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/f/fd/The_effect_of_Rs_and_Gs_in_cAMP_signal_pathway.jpg 2x" data-file-width="418" data-file-height="291" /></a><figcaption>The effect of Rs and Gs in cAMP signal pathway</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg/250px-The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg" decoding="async" width="250" height="181" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg/375px-The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/a/a3/The_effect_of_Ri_and_Gi_in_cAMP_signal_pathway.jpg 2x" data-file-width="394" data-file-height="286" /></a><figcaption>The effect of Ri and Gi in cAMP signal pathway</figcaption></figure> <p>There are two principal signal transduction pathways involving the <a href="/wiki/G_protein-coupled_receptors" class="mw-redirect" title="G protein-coupled receptors">G protein-linked receptors</a>: the <a href="/wiki/Cyclic_adenosine_monophosphate" title="Cyclic adenosine monophosphate">cAMP</a> signal pathway and the <a href="/wiki/Phosphatidylinositol" title="Phosphatidylinositol">phosphatidylinositol</a> signal pathway.<sup id="cite_ref-Gilman_1987_5-1" class="reference"><a href="#cite_note-Gilman_1987-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="cAMP_signal_pathway">cAMP signal pathway</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=19" title="Edit section: cAMP signal pathway"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP-dependent pathway</a></div> <p>The cAMP signal transduction contains five main characters: stimulative <a href="/wiki/Hormone" title="Hormone">hormone</a> receptor (Rs) or inhibitory <a href="/wiki/Hormone_receptor" title="Hormone receptor">hormone receptor</a> (Ri); stimulative regulative G-protein (Gs) or inhibitory regulative G-protein (Gi); <a href="/wiki/Adenylyl_cyclase" title="Adenylyl cyclase">adenylyl cyclase</a>; <a href="/wiki/Protein_kinase_A" title="Protein kinase A">protein kinase A</a> (PKA); and cAMP <a href="/wiki/Phosphodiesterase" title="Phosphodiesterase">phosphodiesterase</a>. </p><p>Stimulative hormone receptor (Rs) is a receptor that can bind with stimulative signal molecules, while inhibitory hormone receptor (Ri) is a receptor that can bind with inhibitory signal molecules. </p><p>Stimulative regulative G-protein is a G-protein linked to stimulative hormone receptor (Rs), and its α subunit upon activation could stimulate the activity of an enzyme or other intracellular metabolism. On the contrary, inhibitory regulative G-protein is linked to an inhibitory hormone receptor, and its α subunit upon activation could inhibit the activity of an enzyme or other intracellular metabolism. </p><p>Adenylyl cyclase is a 12-transmembrane glycoprotein that catalyzes the conversion of ATP to cAMP with the help of cofactor Mg<sup>2+</sup> or Mn<sup>2+</sup>. The cAMP produced is a second messenger in cellular metabolism and is an allosteric activator of protein kinase A. </p><p>Protein kinase A is an important enzyme in cell metabolism due to its ability to regulate cell metabolism by phosphorylating specific committed enzymes in the metabolic pathway. It can also regulate specific gene expression, cellular secretion, and membrane permeability. The protein enzyme contains two catalytic subunits and two regulatory subunits. When there is no cAMP，the complex is inactive. When cAMP binds to the regulatory subunits, their conformation is altered, causing the dissociation of the regulatory subunits, which activates protein kinase A and allows further biological effects. </p><p>These signals then can be terminated by cAMP phosphodiesterase, which is an enzyme that degrades cAMP to 5'-AMP and inactivates protein kinase A. </p> <div class="mw-heading mw-heading3"><h3 id="Phosphatidylinositol_signal_pathway">Phosphatidylinositol signal pathway</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=20" title="Edit section: Phosphatidylinositol signal pathway"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/IP3/DAG_pathway" class="mw-redirect" title="IP3/DAG pathway">IP3/DAG pathway</a></div> <p>In the <a href="/wiki/Phosphatidylinositol" title="Phosphatidylinositol">phosphatidylinositol</a> signal pathway, the extracellular signal molecule binds with the G-protein receptor (G<sub>q</sub>) on the cell surface and activates <a href="/wiki/Phospholipase_C" title="Phospholipase C">phospholipase C</a>, which is located on the <a href="/wiki/Cell_membrane" title="Cell membrane">plasma membrane</a>. The <a href="/wiki/Lipase" title="Lipase">lipase</a> hydrolyzes <a href="/wiki/Phosphatidylinositol_4,5-bisphosphate" title="Phosphatidylinositol 4,5-bisphosphate">phosphatidylinositol 4,5-bisphosphate</a> (PIP2) into two second messengers: <a href="/wiki/Inositol_trisphosphate" title="Inositol trisphosphate">inositol 1,4,5-trisphosphate (IP3)</a> and <a href="/wiki/Diacylglycerol" class="mw-redirect" title="Diacylglycerol">diacylglycerol</a> (DAG). IP3 binds with the <a href="/wiki/IP3_receptor" class="mw-redirect" title="IP3 receptor">IP3 receptor</a> in the membrane of the smooth endoplasmic reticulum and mitochondria to open Ca<sup>2+</sup> channels. DAG helps activate <a href="/wiki/Protein_kinase_C" title="Protein kinase C">protein kinase C</a> (PKC), which phosphorylates many other proteins, changing their catalytic activities, leading to cellular responses. </p><p>The effects of Ca<sup>2+</sup> are also remarkable: it cooperates with DAG in activating PKC and can activate the <a href="/wiki/Ca2%2B/calmodulin-dependent_protein_kinase" class="mw-redirect" title="Ca2+/calmodulin-dependent protein kinase">CaM kinase</a> pathway, in which calcium-modulated protein <a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a> (CaM) binds Ca<sup>2+</sup>, undergoes a change in conformation, and activates CaM kinase II, which has unique ability to increase its binding affinity to CaM by autophosphorylation, making CaM unavailable for the activation of other enzymes. The kinase then phosphorylates target enzymes, regulating their activities. The two signal pathways are connected together by Ca<sup>2+</sup>-CaM, which is also a regulatory subunit of adenylyl cyclase and phosphodiesterase in the cAMP signal pathway. </p> <div class="mw-heading mw-heading2"><h2 id="Receptor_regulation">Receptor regulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=21" title="Edit section: Receptor regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>GPCRs become desensitized when exposed to their ligand for a long period of time. There are two recognized forms of desensitization: 1) <a href="/wiki/Homologous_desensitization" title="Homologous desensitization">homologous desensitization</a>, in which the activated GPCR is downregulated; and 2) <a href="/wiki/Heterologous_desensitization" class="mw-redirect" title="Heterologous desensitization">heterologous desensitization</a>, wherein the activated GPCR causes downregulation of a different GPCR. The key reaction of this downregulation is the <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a> of the intracellular (or <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasmic</a>) receptor domain by <a href="/wiki/Protein_kinase" title="Protein kinase">protein kinases</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Phosphorylation_by_cAMP-dependent_protein_kinases">Phosphorylation by cAMP-dependent protein kinases</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=22" title="Edit section: Phosphorylation by cAMP-dependent protein kinases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cyclic AMP-dependent protein kinases (<a href="/wiki/Protein_kinase_A" title="Protein kinase A">protein kinase A</a>) are activated by the signal chain coming from the G protein (that was activated by the receptor) via <a href="/wiki/Adenylate_cyclase" class="mw-redirect" title="Adenylate cyclase">adenylate cyclase</a> and <a href="/wiki/Cyclic_AMP" class="mw-redirect" title="Cyclic AMP">cyclic AMP</a> (cAMP). In a <i>feedback mechanism</i>, these activated kinases phosphorylate the receptor. The longer the receptor remains active the more kinases are activated and the more receptors are phosphorylated. In <a href="/wiki/Beta-2_adrenergic_receptor" title="Beta-2 adrenergic receptor">β<sub>2</sub>-adrenoceptors</a>, this phosphorylation results in the switching of the coupling from the G<sub>s</sub> class of G-protein to the <a href="/wiki/Gi_alpha_subunit" title="Gi alpha subunit">G<sub>i</sub></a> class.<sup id="cite_ref-pmid11053129_52-0" class="reference"><a href="#cite_note-pmid11053129-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> cAMP-dependent PKA mediated phosphorylation can cause heterologous desensitisation in receptors other than those activated.<sup id="cite_ref-pmid14744258_53-0" class="reference"><a href="#cite_note-pmid14744258-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Phosphorylation_by_GRKs">Phosphorylation by GRKs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=23" title="Edit section: Phosphorylation by GRKs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/G_protein-coupled_receptor_kinases" class="mw-redirect" title="G protein-coupled receptor kinases">G protein-coupled receptor kinases</a> (GRKs) are protein kinases that phosphorylate only active GPCRs.<sup id="cite_ref-Trimarco2013_54-0" class="reference"><a href="#cite_note-Trimarco2013-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> G-protein-coupled receptor kinases (GRKs) are key modulators of G-protein-coupled receptor (GPCR) signaling. They constitute a family of seven mammalian serine-threonine protein kinases that phosphorylate agonist-bound receptor. GRKs-mediated receptor phosphorylation rapidly initiates profound impairment of receptor signaling and desensitization. Activity of GRKs and subcellular targeting is tightly regulated by interaction with receptor domains, G protein subunits, lipids, anchoring proteins and calcium-sensitive proteins.<sup id="cite_ref-pmid14499340_55-0" class="reference"><a href="#cite_note-pmid14499340-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>Phosphorylation of the receptor can have two consequences: </p> <ol><li><i>Translocation</i>: The receptor is, along with the part of the membrane it is embedded in, brought to the inside of the cell, where it is dephosphorylated within the acidic vesicular environment<sup id="cite_ref-pmid8995214_56-0" class="reference"><a href="#cite_note-pmid8995214-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> and then brought back. This mechanism is used to regulate long-term exposure, for example, to a hormone, by allowing resensitisation to follow desensitisation. Alternatively, the receptor may undergo lysozomal degradation, or remain internalised, where it is thought to participate in the initiation of signalling events, the nature of which depending on the internalised vesicle's subcellular localisation.<sup id="cite_ref-pmid14744258_53-1" class="reference"><a href="#cite_note-pmid14744258-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup></li> <li><i><a href="/wiki/Arrestin" title="Arrestin">Arrestin</a> linking</i>: The phosphorylated receptor can be linked to <i>arrestin</i> molecules that prevent it from binding (and activating) G proteins, in effect switching it off for a short period of time. This mechanism is used, for example, with <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a> in <a href="/wiki/Retina" title="Retina">retina</a> cells to compensate for exposure to bright light. In many cases, arrestin's binding to the receptor is a prerequisite for translocation. For example, beta-arrestin bound to β<sub>2</sub>-adrenoreceptors acts as an adaptor for binding with clathrin, and with the beta-subunit of AP2 (clathrin adaptor molecules); thus, the arrestin here acts as a scaffold assembling the components needed for clathrin-mediated endocytosis of β<sub>2</sub>-adrenoreceptors.<sup id="cite_ref-pmid10770944_57-0" class="reference"><a href="#cite_note-pmid10770944-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10097102_58-0" class="reference"><a href="#cite_note-pmid10097102-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup></li></ol> <div class="mw-heading mw-heading3"><h3 id="Mechanisms_of_GPCR_signal_termination">Mechanisms of GPCR signal termination</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=24" title="Edit section: Mechanisms of GPCR signal termination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As mentioned above, G-proteins may terminate their own activation due to their intrinsic <a href="/wiki/GTPase" title="GTPase">GTP→GDP hydrolysis</a> capability. However, this reaction proceeds at a slow <a href="/wiki/Rate_constant" class="mw-redirect" title="Rate constant">rate</a> (≈0.02 times/sec) and, thus, it would take around 50 seconds for any single G-protein to deactivate if other factors did not come into play. Indeed, there are around 30 <a href="/wiki/Protein_isoform" title="Protein isoform">isoforms</a> of <a href="/wiki/Regulator_of_G_protein_signaling" title="Regulator of G protein signaling">RGS proteins</a> that, when bound to Gα through their <a href="/wiki/GTPase_activating_protein" class="mw-redirect" title="GTPase activating protein">GAP domain</a>, accelerate the hydrolysis rate to ≈30 times/sec. This 1500-fold increase in rate allows for the cell to respond to external signals with high speed, as well as spatial <a href="/wiki/Angular_resolution" title="Angular resolution">resolution</a> due to limited amount of <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">second messenger</a> that can be generated and limited distance a G-protein can diffuse in 0.03 seconds. For the most part, the RGS proteins are <a href="/wiki/Promiscuous" class="mw-redirect" title="Promiscuous">promiscuous</a> in their ability to deactivate G-proteins, while which RGS is involved in a given signaling pathway seems more determined by the tissue and GPCR involved than anything else. In addition, RGS proteins have the additional function of increasing the rate of GTP-GDP exchange at GPCRs, (i.e., as a sort of co-GEF) further contributing to the time resolution of GPCR signaling. </p><p>In addition, the GPCR may be <a href="/wiki/Homologous_desensitization" title="Homologous desensitization">desensitized</a> itself. This can occur as: </p> <ol><li>a direct result of <a href="/wiki/Receptor_theory" title="Receptor theory">ligand occupation</a>, wherein the change in <a href="/wiki/Protein_conformation" class="mw-redirect" title="Protein conformation">conformation</a> allows recruitment of <a href="/wiki/G_protein-coupled_receptor_kinase" title="G protein-coupled receptor kinase">GPCR-Regulating Kinases</a> (GRKs), which go on to <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylate</a> various <a href="/wiki/Serine" title="Serine">serine</a>/<a href="/wiki/Threonine" title="Threonine">threonine</a> residues of IL-3 and the <a href="/wiki/C-terminal" class="mw-redirect" title="C-terminal">C-terminal</a> tail. Upon GRK phosphorylation, the GPCR's affinity for <a href="/wiki/Arrestin" title="Arrestin">β-arrestin</a> (β-arrestin-1/2 in most tissues) is increased, at which point β-arrestin may bind and act to both <a href="/wiki/Sterically" class="mw-redirect" title="Sterically">sterically</a> hinder G-protein coupling as well as initiate the process of <a href="/wiki/Receptor-mediated_endocytosis" title="Receptor-mediated endocytosis">receptor internalization</a> through <a href="/wiki/Clathrin-mediated_endocytosis" class="mw-redirect" title="Clathrin-mediated endocytosis">clathrin-mediated endocytosis</a>. Because only the liganded receptor is desensitized by this mechanism, it is called <a href="/wiki/Homologous_desensitization" title="Homologous desensitization">homologous desensitization</a></li> <li>the affinity for β-arrestin may be increased in a ligand occupation and GRK-independent manner through phosphorylation of different ser/thr sites (but also of IL-3 and the C-terminal tail) by PKC and PKA. These phosphorylations are often sufficient to impair G-protein coupling on their own as well.<sup id="cite_ref-pmid18193069_59-0" class="reference"><a href="#cite_note-pmid18193069-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup></li> <li>PKC/PKA may, instead, phosphorylate GRKs, which can also lead to GPCR phosphorylation and β-arrestin binding in an occupation-independent manner. These latter two mechanisms allow for desensitization of one GPCR due to the activities of others, or <a href="/wiki/Heterologous_desensitization" class="mw-redirect" title="Heterologous desensitization">heterologous desensitization</a>. GRKs may also have GAP domains and so may contribute to inactivation through non-<a href="/wiki/Kinase" title="Kinase">kinase</a> mechanisms as well. A combination of these mechanisms may also occur.</li></ol> <p>Once β-arrestin is bound to a GPCR, it undergoes a conformational change allowing it to serve as a scaffolding protein for an adaptor complex termed <a href="/wiki/AP2_adaptors" class="mw-redirect" title="AP2 adaptors">AP-2</a>, which in turn recruits another protein called <a href="/wiki/Clathrin" title="Clathrin">clathrin</a>. If enough receptors in the local area recruit clathrin in this manner, they aggregate and the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">membrane</a> buds inwardly as a result of interactions between the molecules of clathrin, in a process called <a href="/wiki/Opsonization" class="mw-redirect" title="Opsonization">opsonization</a>. Once the pit has been pinched off the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> due to the actions of two other proteins called <a href="/wiki/Amphiphysin" title="Amphiphysin">amphiphysin</a> and <a href="/wiki/Dynamin" title="Dynamin">dynamin</a>, it is now an <a href="/wiki/Endocytosis" title="Endocytosis">endocytic</a> <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicle</a>. At this point, the adapter molecules and clathrin have <a href="/wiki/Dissociated" class="mw-redirect" title="Dissociated">dissociated</a>, and the receptor is either <a href="/wiki/Protein_targeting" title="Protein targeting">trafficked</a> back to the plasma membrane or targeted to <a href="/wiki/Lysosome" title="Lysosome">lysosomes</a> for <a href="/wiki/Proteolysis" title="Proteolysis">degradation</a>. </p><p>At any point in this process, the β-arrestins may also recruit other proteins—such as the <a href="/wiki/Non-receptor_tyrosine_kinase" title="Non-receptor tyrosine kinase">non-receptor tyrosine kinase</a> (nRTK), <a href="/wiki/Src_(gene)" class="mw-redirect" title="Src (gene)">c-SRC</a>—which may activate <a href="/wiki/Extracellular_signal-regulated_kinases" title="Extracellular signal-regulated kinases">ERK1/2</a>, or other <a href="/wiki/Mitogen-activated_protein_kinase" title="Mitogen-activated protein kinase">mitogen-activated protein kinase</a> (MAPK) signaling through, for example, phosphorylation of the <a href="/wiki/Small_GTPase" title="Small GTPase">small GTPase</a>, <a href="/wiki/Ras_subfamily" class="mw-redirect" title="Ras subfamily">Ras</a>, or recruit the proteins of the <a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">ERK cascade</a> directly (i.e., <a href="/wiki/Raf-1" class="mw-redirect" title="Raf-1">Raf-1</a>, <a href="/wiki/Mitogen-activated_protein_kinase_kinase" title="Mitogen-activated protein kinase kinase">MEK</a>, ERK-1/2) at which point signaling is initiated due to their close proximity to one another. Another target of c-SRC are the dynamin molecules involved in endocytosis. Dynamins <a href="/wiki/Polymerization" title="Polymerization">polymerize</a> around the neck of an incoming vesicle, and their phosphorylation by c-SRC provides the energy necessary for the conformational change allowing the final "pinching off" from the membrane. </p> <div class="mw-heading mw-heading3"><h3 id="GPCR_cellular_regulation">GPCR cellular regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=25" title="Edit section: GPCR cellular regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Receptor desensitization is mediated through a combination phosphorylation, β-arr binding, and endocytosis as described above. Downregulation occurs when endocytosed receptor is embedded in an endosome that is trafficked to merge with an organelle called a lysosome. Because lysosomal membranes are rich in proton pumps, their interiors have low pH (≈4.8 vs. the pH≈7.2 cytosol), which acts to denature the GPCRs. In addition, lysosomes contain many <a href="/wiki/Degradative_enzyme" title="Degradative enzyme">degradative enzymes</a>, including proteases, which can function only at such low pH, and so the peptide bonds joining the residues of the GPCR together may be cleaved. Whether or not a given receptor is trafficked to a lysosome, detained in endosomes, or trafficked back to the plasma membrane depends on a variety of factors, including receptor type and magnitude of the signal. GPCR regulation is additionally mediated by gene transcription factors. These factors can increase or decrease gene transcription and thus increase or decrease the generation of new receptors (up- or down-regulation) that travel to the cell membrane. </p> <div class="mw-heading mw-heading2"><h2 id="Receptor_oligomerization">Receptor oligomerization</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=26" title="Edit section: Receptor oligomerization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/GPCR_oligomer" title="GPCR oligomer">GPCR oligomer</a></div> <p>G-protein-coupled receptor oligomerisation is a widespread phenomenon. One of the best-studied examples is the metabotropic <a href="/wiki/GABAB_receptor" title="GABAB receptor">GABA<sub>B</sub> receptor</a>. This so-called constitutive receptor is formed by heterodimerization of <a href="/wiki/GABBR1" title="GABBR1">GABA<sub>B</sub>R1</a> and <a href="/wiki/GABBR2" title="GABBR2">GABA<sub>B</sub>R2</a> subunits. Expression of the GABA<sub>B</sub>R1 without the GABA<sub>B</sub>R2 in heterologous systems leads to retention of the subunit in the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>. Expression of the GABA<sub>B</sub>R2 subunit alone, meanwhile, leads to surface expression of the subunit, although with no functional activity (i.e., the receptor does not bind agonist and cannot initiate a response following exposure to agonist). Expression of the two subunits together leads to plasma membrane expression of functional receptor. It has been shown that GABA<sub>B</sub>R2 binding to GABA<sub>B</sub>R1 causes masking of a retention signal<sup id="cite_ref-pmid10939334_60-0" class="reference"><a href="#cite_note-pmid10939334-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> of functional receptors.<sup id="cite_ref-pmid9872316_61-0" class="reference"><a href="#cite_note-pmid9872316-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Origin_and_diversification_of_the_superfamily">Origin and diversification of the superfamily</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=27" title="Edit section: Origin and diversification of the superfamily"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Signal_transduction" title="Signal transduction">Signal transduction</a> mediated by the superfamily of GPCRs dates back to the origin of <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellularity</a>. Mammalian-like GPCRs are found in <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>, and have been classified according to the <a href="/wiki/GRAFS" title="GRAFS">GRAFS</a> classification system based on GPCR fingerprints.<sup id="cite_ref-pmid22238661_17-1" class="reference"><a href="#cite_note-pmid22238661-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Identification of the superfamily members across the <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> domain, and comparison of the family-specific motifs, have shown that the superfamily of GPCRs have a common origin.<sup id="cite_ref-pmid21402729_62-0" class="reference"><a href="#cite_note-pmid21402729-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Characteristic motifs indicate that three of the five GRAFS families, <i><a href="/wiki/Rhodopsin-like_receptors" title="Rhodopsin-like receptors">Rhodopsin</a></i>, <a href="/wiki/Adhesion_G_protein-coupled_receptor" title="Adhesion G protein-coupled receptor"><i>Adhesion</i></a>, and <i><a href="/wiki/Frizzled" title="Frizzled">Frizzled</a></i>, evolved from the <i><a href="/wiki/Dictyostelium" title="Dictyostelium">Dictyostelium</a> discoideum</i> cAMP receptors before the split of <a href="/wiki/Opisthokont" title="Opisthokont">opisthokonts</a>. Later, the <i><a href="/wiki/Secretin_receptor_family" title="Secretin receptor family">Secretin</a></i> family evolved from the <i>Adhesion</i> GPCR receptor family before the split of <a href="/wiki/Nematode" title="Nematode">nematodes</a>.<sup id="cite_ref-pmid22238661_17-2" class="reference"><a href="#cite_note-pmid22238661-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Insect GPCRs appear to be in their own group and Taste2 is identified as descending from <i>Rhodopsin</i>.<sup id="cite_ref-pmid21402729_62-1" class="reference"><a href="#cite_note-pmid21402729-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Note that the <i>Secretin</i>/<i>Adhesion</i> split is based on presumed function rather than signature, as the classical Class B (7tm_2, <a href="/wiki/Pfam" title="Pfam">Pfam</a> <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00002">PF00002</a>) is used to identify both in the studies. </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=28" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/G_protein-coupled_receptors_database" title="G protein-coupled receptors database">G protein-coupled receptors database</a></li> <li><a href="/wiki/List_of_MeSH_codes_(D12.776)" title="List of MeSH codes (D12.776)">List of MeSH codes (D12.776)</a></li> <li><a href="/wiki/Metabotropic_receptor" title="Metabotropic receptor">Metabotropic receptor</a></li> <li><a href="/wiki/Orphan_receptor" title="Orphan receptor">Orphan receptor</a></li> <li><a href="/wiki/Pepducin" title="Pepducin">Pepducins</a>, a class of drug candidates targeted at GPCRs</li> <li><a href="/wiki/Receptor_activated_solely_by_a_synthetic_ligand" title="Receptor activated solely by a synthetic ligand">Receptor activated solely by a synthetic ligand</a>, a technique for control of cell signaling through synthetic GPCRs</li> <li><a href="/wiki/TOG_superfamily" title="TOG superfamily">TOG superfamily</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=29" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Cherezov_2007-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Cherezov_2007_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Cherezov_2007_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFCherezovRosenbaumHansonRasmussen2007" class="citation journal cs1">Cherezov V, Rosenbaum DM, Hanson MA, Rasmussen SG, Thian FS, Kobilka TS, et al. 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(April 2003). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC153653">"The G protein-coupled receptor repertoires of human and mouse"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>100</b> (8): 4903–8. <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/2003PNAS..100.4903V">2003PNAS..100.4903V</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.0230374100">10.1073/pnas.0230374100</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/PMC153653">153653</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/12679517">12679517</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=The+G+protein-coupled+receptor+repertoires+of+human+and+mouse&rft.volume=100&rft.issue=8&rft.pages=4903-8&rft.date=2003-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC153653%23id-name%3DPMC&rft_id=info%3Apmid%2F12679517&rft_id=info%3Adoi%2F10.1073%2Fpnas.0230374100&rft_id=info%3Abibcode%2F2003PNAS..100.4903V&rft.aulast=Vassilatis&rft.aufirst=DK&rft.au=Hohmann%2C+JG&rft.au=Zeng%2C+H&rft.au=Li%2C+F&rft.au=Ranchalis%2C+JE&rft.au=Mortrud%2C+MT&rft.au=Brown%2C+A&rft.au=Rodriguez%2C+SS&rft.au=Weller%2C+JR&rft.au=Wright%2C+AC&rft.au=Bergmann%2C+JE&rft.au=Gaitanaris%2C+GA&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC153653&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.bio-balance.com/Ref.htm">"GPCR Reference Library"</a><span class="reference-accessdate">. Retrieved <span class="nowrap">11 August</span> 2008</span>. <q>Reference for molecular and mathematical models for the initial receptor response</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=GPCR+Reference+Library&rft_id=https%3A%2F%2Fwww.bio-balance.com%2FRef.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.nobelprize.org/nobel_prizes/chemistry/laureates/2012/popular-chemistryprize2012.pdf">"The Nobel Prize in Chemistry 2012"</a> <span class="cs1-format">(PDF)</span>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20121018052846/http://www.nobelprize.org/nobel_prizes/chemistry/laureates/2012/popular-chemistryprize2012.pdf">Archived</a> <span class="cs1-format">(PDF)</span> from the original on 18 October 2012<span class="reference-accessdate">. Retrieved <span class="nowrap">10 October</span> 2012</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=The+Nobel+Prize+in+Chemistry+2012&rft_id=https%3A%2F%2Fwww.nobelprize.org%2Fnobel_prizes%2Fchemistry%2Flaureates%2F2012%2Fpopular-chemistryprize2012.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=G_protein-coupled_receptor&action=edit&section=31" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:G_protein-coupled_receptors" class="extiw" title="commons:Category:G protein-coupled receptors">G protein-coupled receptors</a></span>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=G-protein-coupled+receptors">G-protein-coupled+receptors</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://www.creative-biogene.com/Product/GPCR-list-11.html">GPCR Cell Line</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20150403202411/https://www.creative-biogene.com/Product/GPCR-list-11.html">Archived</a> 3 April 2015 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.guidetopharmacology.org/GRAC/ReceptorFamiliesForward?type=GPCR">"IUPHAR/BPS Guide to PHARMACOLOGY Database (GPCRs)"</a>. <i>IUPHAR Database</i>. University of Edinburgh / International Union of Basic and Clinical Pharmacology<span class="reference-accessdate">. Retrieved <span class="nowrap">6 February</span> 2019</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=IUPHAR+Database&rft.atitle=IUPHAR%2FBPS+Guide+to+PHARMACOLOGY+Database+%28GPCRs%29&rft_id=https%3A%2F%2Fwww.guidetopharmacology.org%2FGRAC%2FReceptorFamiliesForward%3Ftype%3DGPCR&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://docs.gpcrdb.org/about.html">"GPCRdb"</a>. <q>Data, diagrams and web tools for G protein-coupled receptors (GPCRs).</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=GPCRdb&rft_id=https%3A%2F%2Fdocs.gpcrdb.org%2Fabout.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span>; <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMunkIsbergMordalskiHarpsøe2016" class="citation journal cs1">Munk C, Isberg V, Mordalski S, Harpsøe K, Rataj K, Hauser AS, et al. 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Archived from <a rel="nofollow" class="external text" href="https://www.gproteincoupledreceptors.net/">the original</a> on 23 July 2011<span class="reference-accessdate">. Retrieved <span class="nowrap">10 November</span> 2010</span>. <q>a classification of GPCRs</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=G+Protein-Coupled+Receptors+on+the+NET&rft_id=https%3A%2F%2Fwww.gproteincoupledreceptors.net%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20130725122424/https://gpcr.scripps.edu/">"PSI GPCR Network Center"</a>. Archived from <a rel="nofollow" class="external text" href="https://gpcr.scripps.edu">the original</a> on 25 July 2013<span class="reference-accessdate">. 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(identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26190572">26190572</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Structure&rft.atitle=GPCR-I-TASSER%3A+A+Hybrid+Approach+to+G+Protein-Coupled+Receptor+Structure+Modeling+and+the+Application+to+the+Human+Genome&rft.volume=23&rft.issue=8&rft.pages=1538-1549&rft.date=2015-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4526412%23id-name%3DPMC&rft_id=info%3Apmid%2F26190572&rft_id=info%3Adoi%2F10.1016%2Fj.str.2015.06.007&rft.aulast=Zhang&rft.aufirst=J&rft.au=Yang%2C+J&rft.au=Jang%2C+R&rft.au=Zhang%2C+Y&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4526412&rfr_id=info%3Asid%2Fen.wikipedia.org%3AG+protein-coupled+receptor" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output 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proteins</a>, <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a>: <a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">cell surface receptors</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink selflink">G protein-coupled receptor</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;">Class 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/Eicosanoid_receptor" title="Eicosanoid receptor">Eicosanoid receptor</a> (<a href="/wiki/Prostaglandin_receptor" title="Prostaglandin receptor">Prostaglandin receptor</a>)</li> <li><a href="/wiki/Protease-activated_receptor" title="Protease-activated receptor">Protease-activated receptor</a></li> <li><a href="/wiki/Neurotransmitter_receptor" title="Neurotransmitter receptor">Neurotransmitter receptor</a></li> <li><a href="/wiki/Purinergic_receptor" title="Purinergic receptor">Purinergic receptor</a></li> <li><a href="/wiki/Biogenic_amine_receptor" title="Biogenic amine receptor">Biogenic amine receptor</a></li> <li><a href="/wiki/Olfactory_receptor" title="Olfactory receptor">Olfactory receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite;">Class B</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/Secretin_receptor" title="Secretin receptor">Secretin receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite;">Class C</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/Metabotropic_glutamate_receptor" title="Metabotropic glutamate receptor">Metabotropic glutamate receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite;">Class D</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/Vomeronasal_receptor" title="Vomeronasal receptor">Pheromone receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite;">Class E</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/CAMP_receptor" class="mw-redirect" title="CAMP receptor">cAMP receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite;">Class F</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/Frizzled" title="Frizzled">Frizzled</a>/<a href="/wiki/Smoothened" title="Smoothened">smoothened</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/Ligand-gated_ion_channel" title="Ligand-gated ion channel">Ligand-gated ion channel</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/Purinergic_receptor" title="Purinergic receptor">Purinergic receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Enzyme-linked_receptor" title="Enzyme-linked receptor">Enzyme-linked receptor</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/Serine/threonine-specific_protein_kinase" title="Serine/threonine-specific protein kinase">Serine/threonine-specific protein kinase</a></li> <li><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">Receptor tyrosine kinase</a></li> <li><a href="/wiki/Guanylate_cyclase" title="Guanylate cyclase">Guanylate cyclase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other/ungrouped</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/Asialoglycoprotein_receptor" title="Asialoglycoprotein receptor">Asialoglycoprotein receptor</a></li> <li><a href="/wiki/Tumor_necrosis_factor_receptor" class="mw-redirect" title="Tumor necrosis factor receptor">Tumor necrosis factor receptor</a></li> <li><a href="/wiki/Immunoglobulin_superfamily" title="Immunoglobulin superfamily">Immunoglobulin superfamily</a></li> <li><a href="/wiki/N-Acetylglucosamine_receptor" title="N-Acetylglucosamine receptor"><i>N</i>-Acetylglucosamine receptor</a></li> <li><a href="/wiki/Neuropilin" title="Neuropilin">Neuropilins</a></li> <li><a href="/wiki/Transferrin_receptor" title="Transferrin receptor">Transferrin receptor</a></li> <li><a href="/wiki/EDAR" class="mw-redirect" title="EDAR">EDAR</a></li> <li><a href="/wiki/Lipoprotein_receptor-related_protein" title="Lipoprotein receptor-related protein">Lipoprotein receptor-related protein</a></li> <li><a href="/wiki/Progestin_and_adipoQ_receptor" title="Progestin and adipoQ receptor">Progestin and adipoQ receptor</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="Cell_surface_receptor:_G_protein-coupled_receptors" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed 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:G_protein-coupled_receptors" title="Template:G protein-coupled receptors"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:G_protein-coupled_receptors" title="Template talk:G protein-coupled receptors"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:G_protein-coupled_receptors" title="Special:EditPage/Template:G protein-coupled receptors"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_surface_receptor:_G_protein-coupled_receptors" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">Cell surface receptor</a>: <a class="mw-selflink selflink">G protein-coupled receptors</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Class_A:_Rhodopsin-like" style="font-size:114%;margin:0 4em"><a href="/wiki/Rhodopsin-like_receptors" title="Rhodopsin-like receptors">Class A</a>: <a href="/wiki/Rhodopsin" title="Rhodopsin">Rhodopsin</a>-like</div></th></tr><tr><td colspan="2" class="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:7em;text-align:left;"><a href="/wiki/Neurotransmitter_receptor" title="Neurotransmitter receptor">Neurotransmitter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="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:7em;text-align:left;"><a href="/wiki/Adrenergic_receptor" title="Adrenergic receptor">Adrenergic</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alpha-1_adrenergic_receptor" title="Alpha-1 adrenergic receptor">α1</a> (<a href="/wiki/Alpha-1A_adrenergic_receptor" title="Alpha-1A adrenergic receptor">A</a></li> <li><a href="/wiki/Alpha-1B_adrenergic_receptor" title="Alpha-1B adrenergic receptor">B</a></li> <li><a href="/wiki/Alpha-1D_adrenergic_receptor" title="Alpha-1D adrenergic receptor">D</a>)</li> <li><a href="/wiki/Alpha-2_adrenergic_receptor" title="Alpha-2 adrenergic receptor">α2</a> (<a href="/wiki/Alpha-2A_adrenergic_receptor" title="Alpha-2A adrenergic receptor">A</a></li> <li><a href="/wiki/Alpha-2B_adrenergic_receptor" title="Alpha-2B adrenergic receptor">B</a></li> <li><a href="/wiki/Alpha-2C_adrenergic_receptor" title="Alpha-2C adrenergic receptor">C</a>)</li> <li><a href="/wiki/Beta-1_adrenergic_receptor" title="Beta-1 adrenergic receptor">β1</a></li> <li><a href="/wiki/Beta-2_adrenergic_receptor" title="Beta-2 adrenergic receptor">β2</a></li> <li><a href="/wiki/Beta-3_adrenergic_receptor" title="Beta-3 adrenergic receptor">β3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;"><a href="/wiki/Purinergic_receptor" title="Purinergic receptor">Purinergic</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adenosine_receptor" title="Adenosine receptor">Adenosine</a> (<a href="/wiki/Adenosine_A1_receptor" title="Adenosine A1 receptor">A1</a></li> <li><a href="/wiki/Adenosine_A2A_receptor" title="Adenosine A2A receptor">A2A</a></li> <li><a href="/wiki/Adenosine_A2B_receptor" title="Adenosine A2B receptor">A2B</a></li> <li><a href="/wiki/Adenosine_A3_receptor" title="Adenosine A3 receptor">A3</a>)</li> <li><a href="/wiki/P2Y_receptor" title="P2Y receptor">P2Y</a> (<a href="/wiki/P2RY1" title="P2RY1">1</a></li> <li><a href="/wiki/P2RY2" title="P2RY2">2</a></li> <li><a href="/wiki/P2RY4" title="P2RY4">4</a></li> <li><a href="/wiki/LPAR6" title="LPAR6">5</a></li> <li><a href="/wiki/P2RY6" title="P2RY6">6</a></li> <li><a href="/wiki/P2RY8" title="P2RY8">8</a></li> <li><a href="/wiki/GPR23" class="mw-redirect" title="GPR23">9</a></li> <li><a href="/wiki/P2RY10" title="P2RY10">10</a></li> <li><a href="/wiki/P2RY11" title="P2RY11">11</a></li> <li><a href="/wiki/P2Y12" title="P2Y12">12</a></li> <li><a href="/wiki/P2RY13" title="P2RY13">13</a></li> <li><a href="/wiki/P2RY14" title="P2RY14">14</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;"><a href="/wiki/5-HT_receptor" title="5-HT receptor">Serotonin</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>(all but <a href="/wiki/5-HT3_receptor" title="5-HT3 receptor">5-HT3</a>)</i> <a href="/wiki/5-HT1_receptor" title="5-HT1 receptor">5-HT1</a> (<a href="/wiki/5-HT1A_receptor" title="5-HT1A receptor">A</a></li> <li><a href="/wiki/5-HT1B_receptor" title="5-HT1B receptor">B</a></li> <li><a href="/wiki/5-HT1D_receptor" title="5-HT1D receptor">D</a></li> <li><a href="/wiki/5-HT1E_receptor" title="5-HT1E receptor">E</a></li> <li><a href="/wiki/5-HT1F_receptor" title="5-HT1F receptor">F</a>)</li> <li><a href="/wiki/5-HT2_receptor" title="5-HT2 receptor">5-HT2</a> (<a href="/wiki/5-HT2A_receptor" title="5-HT2A receptor">A</a></li> <li><a href="/wiki/5-HT2B_receptor" title="5-HT2B receptor">B</a></li> <li><a href="/wiki/5-HT2C_receptor" title="5-HT2C receptor">C</a>)</li> <li><i>5-HT</i> (<a href="/wiki/5-HT4_receptor" title="5-HT4 receptor">4</a></li> <li><a href="/wiki/5-HT5A_receptor" title="5-HT5A receptor">5A</a></li> <li><a href="/wiki/5-HT6_receptor" title="5-HT6 receptor">6</a></li> <li><a href="/wiki/5-HT7_receptor" title="5-HT7 receptor">7</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Muscarinic_acetylcholine_receptor" title="Muscarinic acetylcholine receptor">Acetylcholine</a> (<a href="/wiki/Muscarinic_acetylcholine_receptor_M1" title="Muscarinic acetylcholine receptor M1">M1</a></li> <li><a href="/wiki/Muscarinic_acetylcholine_receptor_M2" title="Muscarinic acetylcholine receptor M2">M2</a></li> <li><a href="/wiki/Muscarinic_acetylcholine_receptor_M3" title="Muscarinic acetylcholine receptor M3">M3</a></li> <li><a href="/wiki/Muscarinic_acetylcholine_receptor_M4" title="Muscarinic acetylcholine receptor M4">M4</a></li> <li><a href="/wiki/Muscarinic_acetylcholine_receptor_M5" title="Muscarinic acetylcholine receptor M5">M5</a>)</li> <li><a href="/wiki/Dopamine_receptor" title="Dopamine receptor">Dopamine</a> <ul><li><a href="/wiki/Dopamine_receptor_D1" title="Dopamine receptor D1">D1</a></li> <li><a href="/wiki/Dopamine_receptor_D2" title="Dopamine receptor D2">D2</a></li> <li><a href="/wiki/Dopamine_receptor_D3" title="Dopamine receptor D3">D3</a></li> <li><a href="/wiki/Dopamine_receptor_D4" title="Dopamine receptor D4">D4</a></li> <li><a href="/wiki/Dopamine_receptor_D5" title="Dopamine receptor D5">D5</a></li></ul></li> <li><a href="/wiki/GHB_receptor" title="GHB receptor">GHB receptor</a></li> <li><a href="/wiki/Histamine_receptor" title="Histamine receptor">Histamine</a> <ul><li><a href="/wiki/Histamine_H1_receptor" title="Histamine H1 receptor">H1</a></li> <li><a href="/wiki/Histamine_H2_receptor" title="Histamine H2 receptor">H2</a></li> <li><a href="/wiki/Histamine_H3_receptor" title="Histamine H3 receptor">H3</a></li> <li><a href="/wiki/Histamine_H4_receptor" title="Histamine H4 receptor">H4</a></li></ul></li> <li><a href="/wiki/Melatonin_receptor" title="Melatonin receptor">Melatonin</a> (<a href="/wiki/Melatonin_receptor_1A" title="Melatonin receptor 1A">1A</a></li> <li><a href="/wiki/Melatonin_receptor_1B" title="Melatonin receptor 1B">1B</a></li> <li><a href="/wiki/Melatonin_receptor_1C" title="Melatonin receptor 1C">1C</a>)</li> <li><a href="/wiki/Trace_amine-associated_receptor" title="Trace amine-associated receptor">TAAR</a> (<a href="/wiki/TAAR1" title="TAAR1">1</a></li> <li><a href="/wiki/TAAR2" title="TAAR2">2</a></li> <li><a href="/wiki/TAAR5" title="TAAR5">5</a></li> <li><a href="/wiki/TAAR6" title="TAAR6">6</a></li> <li><a href="/wiki/TAAR8" title="TAAR8">8</a></li> <li><a href="/wiki/TAAR9" title="TAAR9">9</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Metabolites and<br />signaling molecules</th><td class="navbox-list-with-group navbox-list navbox-odd" style="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:7em;text-align:left;"><a href="/wiki/Eicosanoid_receptor" title="Eicosanoid receptor">Eicosanoid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>CysLT</i> (<a href="/wiki/Cysteinyl_leukotriene_receptor_1" title="Cysteinyl leukotriene receptor 1">1</a></li> <li><a href="/wiki/Cysteinyl_leukotriene_receptor_2" title="Cysteinyl leukotriene receptor 2">2</a>)</li> <li><a href="/wiki/Leukotriene_B4" title="Leukotriene B4">LTB4</a> <ul><li><a href="/wiki/Leukotriene_B4_receptor" title="Leukotriene B4 receptor">1</a></li> <li><a href="/wiki/Leukotriene_B4_receptor_2" title="Leukotriene B4 receptor 2">2</a></li></ul></li> <li><a href="/wiki/Formyl_peptide_receptor_2" title="Formyl peptide receptor 2"> FPRL1</a></li> <li><a href="/wiki/Oxoeicosanoid_receptor_1" title="Oxoeicosanoid receptor 1">OXE</a></li> <li><a href="/wiki/Prostaglandin_receptor" title="Prostaglandin receptor">Prostaglandin</a> <ul><li><i>DP</i> (<a href="/wiki/Prostaglandin_D2_receptor" title="Prostaglandin D2 receptor">1</a></li> <li><a href="/wiki/GPR44" class="mw-redirect" title="GPR44">2</a>), <i>EP</i> (<a href="/wiki/EP1_receptor" class="mw-redirect" title="EP1 receptor">1</a></li> <li><a href="/wiki/Prostaglandin_E2_receptor" title="Prostaglandin E2 receptor">2</a></li> <li><a href="/wiki/Prostaglandin_E_receptor_3" class="mw-redirect" title="Prostaglandin E receptor 3">3</a></li> <li><a href="/wiki/EP4_receptor" class="mw-redirect" title="EP4 receptor">4</a>), <a href="/wiki/Prostaglandin_F_receptor" title="Prostaglandin F receptor">FP</a></li></ul></li> <li><a href="/wiki/Prostacyclin_receptor" title="Prostacyclin receptor">Prostacyclin</a></li> <li><a href="/wiki/Thromboxane_receptor" title="Thromboxane receptor">Thromboxane</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/G_protein-coupled_bile_acid_receptor" title="G protein-coupled bile acid receptor">Bile acid</a></li> <li><a href="/wiki/Cannabinoid_receptor" title="Cannabinoid receptor">Cannabinoid</a> (<a href="/wiki/Cannabinoid_receptor_type_1" class="mw-redirect" title="Cannabinoid receptor type 1">CB1</a></li> <li><a href="/wiki/Cannabinoid_receptor_type_2" class="mw-redirect" title="Cannabinoid receptor type 2">CB2</a>, <i>GPR</i> (<a href="/wiki/GPR18" class="mw-redirect" title="GPR18">18</a></li> <li><a href="/wiki/GPR55" title="GPR55">55</a></li> <li><a href="/wiki/GPR119" title="GPR119">119</a>))</li> <li><a href="/wiki/GPR183" title="GPR183">EBI2</a></li> <li><a href="/wiki/GPER" title="GPER">Estrogen</a></li> <li><a href="/wiki/Free_fatty_acid_receptor" title="Free fatty acid receptor">Free fatty acid</a> (<a href="/wiki/Free_fatty_acid_receptor_1" title="Free fatty acid receptor 1">1</a></li> <li><a href="/wiki/Free_fatty_acid_receptor_2" title="Free fatty acid receptor 2">2</a></li> <li><a href="/wiki/Free_fatty_acid_receptor_3" title="Free fatty acid receptor 3">3</a></li> <li><a href="/wiki/GPR42" title="GPR42">4</a>)</li> <li><a href="/wiki/Hydroxycarboxylic_acid_receptor" title="Hydroxycarboxylic acid receptor">Hydroxycarboxylic acids</a> <ul><li><a href="/wiki/Hydroxycarboxylic_acid_receptor_1" title="Hydroxycarboxylic acid receptor 1">1</a></li> <li><a href="/wiki/Hydroxycarboxylic_acid_receptor_2" title="Hydroxycarboxylic acid receptor 2">2</a></li> <li><a href="/wiki/Hydroxycarboxylic_acid_receptor_3" title="Hydroxycarboxylic acid receptor 3">3</a></li></ul></li> <li><a href="/wiki/Lysophosphatidic_acid" title="Lysophosphatidic acid">Lysophosphatidic acid</a> (<a href="/wiki/LPAR1" title="LPAR1">1</a></li> <li><a href="/wiki/LPAR2" title="LPAR2">2</a></li> <li><a href="/wiki/LPAR3" title="LPAR3">3</a></li> <li><a href="/wiki/LPAR4" title="LPAR4">4</a></li> <li><a href="/wiki/LPAR5" title="LPAR5">5</a></li> <li><a href="/wiki/LPAR6" title="LPAR6">6</a>)</li> <li><a href="/wiki/Lysophospholipid_receptor" title="Lysophospholipid receptor">Lysophospholipid</a> (<a href="/wiki/S1PR1" title="S1PR1">1</a></li> <li><a href="/wiki/LPAR1" title="LPAR1">2</a></li> <li><a href="/wiki/S1PR3" title="S1PR3">3</a></li> <li><a href="/wiki/LPAR2" title="LPAR2">4</a></li> <li><a href="/wiki/S1PR2" title="S1PR2">5</a></li> <li><a href="/wiki/S1PR4" title="S1PR4">6</a></li> <li><a href="/wiki/LPAR3" title="LPAR3">7</a></li> <li><a href="/wiki/S1PR5" title="S1PR5">8</a>)</li> <li><a href="/wiki/OXGR1" title="OXGR1">Oxoglutarate</a></li> <li><a href="/wiki/Platelet-activating_factor_receptor" title="Platelet-activating factor receptor">PAF</a></li> <li><a href="/wiki/Sphingosine-1-phosphate" title="Sphingosine-1-phosphate">Sphingosine-1-phosphate</a> (<a href="/wiki/S1PR1" title="S1PR1">1</a></li> <li><a href="/wiki/S1PR2" title="S1PR2">2</a></li> <li><a href="/wiki/S1PR3" title="S1PR3">3</a></li> <li><a href="/wiki/S1PR4" title="S1PR4">4</a></li> <li><a href="/wiki/S1PR5" title="S1PR5">5</a>)</li> <li><a href="/wiki/SUCNR1" title="SUCNR1">Succinate</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Peptide</th><td class="navbox-list-with-group navbox-list navbox-odd" style="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:7em;text-align:left;"><a href="/wiki/GPCR_neuropeptide_receptor" title="GPCR neuropeptide receptor">Neuropeptide</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Neuropeptide_B/W_receptor" title="Neuropeptide B/W receptor">B/W</a> (<a href="/wiki/Neuropeptides_B/W_receptor_1" title="Neuropeptides B/W receptor 1">1</a></li> <li><a href="/wiki/Neuropeptides_B/W_receptor_2" title="Neuropeptides B/W receptor 2">2</a>)</li> <li><a href="/wiki/Neuropeptide_FF_receptor" title="Neuropeptide FF receptor">FF</a> (<a href="/wiki/Neuropeptide_FF_receptor_1" title="Neuropeptide FF receptor 1">1</a></li> <li><a href="/wiki/Neuropeptide_FF_receptor_2" title="Neuropeptide FF receptor 2">2</a>)</li> <li><a href="/wiki/Neuropeptide_S_receptor" title="Neuropeptide S receptor">S</a></li> <li><a href="/wiki/Neuropeptide_Y_receptor" title="Neuropeptide Y receptor">Y</a> (<a href="/wiki/Neuropeptide_Y_receptor_Y1" title="Neuropeptide Y receptor Y1">1</a></li> <li><a href="/wiki/Neuropeptide_Y_receptor_Y2" title="Neuropeptide Y receptor Y2">2</a></li> <li><a href="/wiki/Pancreatic_polypeptide_receptor_1" title="Pancreatic polypeptide receptor 1">4</a></li> <li><a href="/wiki/Neuropeptide_Y_receptor_Y5" title="Neuropeptide Y receptor Y5">5</a>)</li> <li><i>Neuromedin</i> (<a href="/wiki/Neuromedin_B_receptor" title="Neuromedin B receptor">B</a></li> <li><a href="/wiki/Neuromedin_U_receptor" title="Neuromedin U receptor">U</a> (<a href="/wiki/Neuromedin_U_receptor_1" title="Neuromedin U receptor 1">1</a></li> <li><a href="/wiki/Neuromedin_U_receptor_2" title="Neuromedin U receptor 2">2</a>))</li> <li><a href="/wiki/Neurotensin_receptor" title="Neurotensin receptor">Neurotensin</a> (<a href="/wiki/Neurotensin_receptor_1" title="Neurotensin receptor 1">1</a></li> <li><a href="/wiki/Neurotensin_receptor_2" title="Neurotensin receptor 2">2</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anaphylatoxin_receptors" title="Anaphylatoxin receptors">Anaphylatoxin</a> (<a href="/wiki/C3a_receptor" title="C3a receptor">C3a</a></li> <li>C5a (<a href="/wiki/C5a_receptor" title="C5a receptor">1</a></li> <li><a href="/wiki/C5AR2" title="C5AR2">2</a>))</li> <li><a href="/wiki/Angiotensin_receptor" class="mw-redirect" title="Angiotensin receptor">Angiotensin</a> (<a href="/wiki/Angiotensin_II_receptor_type_1" title="Angiotensin II receptor type 1">1</a></li> <li><a href="/wiki/Angiotensin_II_receptor_type_2" title="Angiotensin II receptor type 2">2</a>)</li> <li><a href="/wiki/Apelin_receptor" title="Apelin receptor">Apelin</a></li> <li><a href="/wiki/Bombesin_receptor" title="Bombesin receptor">Bombesin</a> <ul><li><a href="/wiki/Bombesin-like_receptor_3" title="Bombesin-like receptor 3">BRS3</a></li> <li><a href="/wiki/Gastrin-releasing_peptide_receptor" title="Gastrin-releasing peptide receptor">GRPR</a></li> <li><a href="/wiki/Neuromedin_B_receptor" title="Neuromedin B receptor">NMBR</a>)</li> <li><a href="/wiki/Bradykinin_receptor" title="Bradykinin receptor">Bradykinin</a> (<a href="/wiki/Bradykinin_receptor_B1" title="Bradykinin receptor B1">B1</a></li> <li><a href="/wiki/Bradykinin_receptor_B2" title="Bradykinin receptor B2">B2</a>)</li></ul></li> <li><a href="/wiki/Chemokine_receptor" title="Chemokine receptor">Chemokine</a></li> <li><a href="/wiki/Cholecystokinin_receptor" title="Cholecystokinin receptor">Cholecystokinin</a> (<a href="/wiki/Cholecystokinin_A_receptor" title="Cholecystokinin A receptor">A</a></li> <li><a href="/wiki/Cholecystokinin_B_receptor" title="Cholecystokinin B receptor">B</a>)</li> <li><a href="/wiki/Endothelin_receptor" title="Endothelin receptor">Endothelin</a> <ul><li><a href="/wiki/Endothelin_A_receptor" title="Endothelin A receptor">A</a></li> <li><a href="/wiki/Endothelin_B_receptor" class="mw-redirect" title="Endothelin B receptor">B</a></li></ul></li> <li><a href="/wiki/Formyl_peptide_receptor" title="Formyl peptide receptor">Formyl peptide</a> (<a href="/wiki/Formyl_peptide_receptor_1" title="Formyl peptide receptor 1">1</a></li> <li><a href="/wiki/Formyl_peptide_receptor_2" title="Formyl peptide receptor 2">2</a></li> <li><a href="/wiki/Formyl_peptide_receptor_3" title="Formyl peptide receptor 3">3</a>)</li> <li><a href="/wiki/Follicle-stimulating_hormone_receptor" title="Follicle-stimulating hormone receptor">FSH</a></li> <li><a href="/wiki/Galanin_receptor" title="Galanin receptor">Galanin</a> (<a href="/wiki/Galanin_receptor_1" title="Galanin receptor 1">1</a></li> <li><a href="/wiki/Galanin_receptor_2" title="Galanin receptor 2">2</a></li> <li><a href="/wiki/Galanin_receptor_3" title="Galanin receptor 3">3</a>)</li> <li><a href="/wiki/Gonadotropin-releasing_hormone_receptor" title="Gonadotropin-releasing hormone receptor">Gonadotropin-releasing hormone</a> (<a href="/wiki/GNRHR" title="GNRHR">1</a></li> <li><a href="/wiki/GNRHR2" title="GNRHR2">2</a>)</li> <li><a href="/wiki/Growth_hormone_secretagogue_receptor" title="Growth hormone secretagogue receptor">Ghrelin</a></li> <li><a href="/wiki/KiSS1-derived_peptide_receptor" title="KiSS1-derived peptide receptor"> Kisspeptin</a></li> <li><a href="/wiki/Luteinizing_hormone/choriogonadotropin_receptor" title="Luteinizing hormone/choriogonadotropin receptor">Luteinizing hormone/choriogonadotropin</a></li> <li><a href="/wiki/MAS1_oncogene" title="MAS1 oncogene">MAS</a> (<a href="/wiki/MAS1" title="MAS1">1</a></li> <li><a href="/wiki/MAS1L" title="MAS1L">1L</a></li> <li><a href="/wiki/MRGPRD" title="MRGPRD">D</a></li> <li><a href="/wiki/MRGPRE" title="MRGPRE">E</a></li> <li><a href="/wiki/MRGPRF" title="MRGPRF">F</a></li> <li><a href="/wiki/MRGPRG" title="MRGPRG">G</a></li> <li><a href="/wiki/MRGPRX1" title="MRGPRX1">X1</a></li> <li><a href="/wiki/MRGPRX2" title="MRGPRX2">X2</a></li> <li><a href="/wiki/MRGPRX3" title="MRGPRX3">X3</a></li> <li><a href="/wiki/MRGPRX4" title="MRGPRX4">X4</a>)</li> <li><a href="/wiki/Melanocortin_receptor" title="Melanocortin receptor">Melanocortin</a> (<a href="/wiki/Melanocortin_1_receptor" title="Melanocortin 1 receptor">1</a></li> <li><a href="/wiki/ACTH_receptor" title="ACTH receptor">2</a></li> <li><a href="/wiki/Melanocortin_3_receptor" title="Melanocortin 3 receptor">3</a></li> <li><a href="/wiki/Melanocortin_4_receptor" title="Melanocortin 4 receptor">4</a></li> <li><a href="/wiki/Melanocortin_5_receptor" title="Melanocortin 5 receptor">5</a>)</li> <li><a href="/wiki/Melanin-concentrating_hormone_receptor" title="Melanin-concentrating hormone receptor">MCHR</a> (<a href="/wiki/Melanin-concentrating_hormone_receptor_1" title="Melanin-concentrating hormone receptor 1">1</a></li> <li><a href="/wiki/Melanin-concentrating_hormone_receptor_2" title="Melanin-concentrating hormone receptor 2">2</a>)</li> <li><a href="/wiki/Motilin_receptor" title="Motilin receptor">Motilin</a></li> <li><a href="/wiki/Opioid_receptor" title="Opioid receptor">Opioid</a> (<a href="/wiki/%CE%94-opioid_receptor" title="Δ-opioid receptor">Delta</a></li> <li><a href="/wiki/%CE%9A-opioid_receptor" title="Κ-opioid receptor">Kappa</a></li> <li><a href="/wiki/%CE%9C-opioid_receptor" title="Μ-opioid receptor">Mu</a></li> <li><a href="/wiki/Nociceptin_receptor" title="Nociceptin receptor">Nociceptin</a> & <a href="/wiki/OGFr" title="OGFr">Zeta</a>, but not <a href="/wiki/Sigma_receptor" title="Sigma receptor">Sigma</a>)</li> <li><a href="/wiki/Orexin_receptor" title="Orexin receptor">Orexin</a> (<a href="/wiki/Hypocretin_(orexin)_receptor_1" title="Hypocretin (orexin) receptor 1">1</a></li> <li><a href="/wiki/Hypocretin_(orexin)_receptor_2" title="Hypocretin (orexin) receptor 2">2</a>)</li> <li><a href="/wiki/Oxytocin_receptor" title="Oxytocin receptor">Oxytocin</a></li> <li><a href="/wiki/Prokineticin_receptor" title="Prokineticin receptor">Prokineticin</a> (<a href="/wiki/Prokineticin_receptor_1" title="Prokineticin receptor 1">1</a></li> <li><a href="/wiki/Prokineticin_receptor_2" title="Prokineticin receptor 2">2</a>)</li> <li><a href="/wiki/Prolactin-releasing_peptide_receptor" title="Prolactin-releasing peptide receptor">Prolactin-releasing peptide</a></li> <li><a href="/wiki/Relaxin_receptor" title="Relaxin receptor">Relaxin</a> (<a href="/wiki/Relaxin/insulin-like_family_peptide_receptor_1" title="Relaxin/insulin-like family peptide receptor 1">1</a></li> <li><a href="/wiki/Relaxin/insulin-like_family_peptide_receptor_2" title="Relaxin/insulin-like family peptide receptor 2">2</a></li> <li><a href="/wiki/Relaxin/insulin-like_family_peptide_receptor_3" title="Relaxin/insulin-like family peptide receptor 3">3</a></li> <li><a href="/wiki/Relaxin/insulin-like_family_peptide_receptor_4" title="Relaxin/insulin-like family peptide receptor 4">4</a>)</li> <li><a href="/wiki/Somatostatin_receptor" title="Somatostatin receptor">Somatostatin</a> (<a href="/wiki/Somatostatin_receptor_1" title="Somatostatin receptor 1">1</a></li> <li><a href="/wiki/Somatostatin_receptor_2" title="Somatostatin receptor 2">2</a></li> <li><a href="/wiki/Somatostatin_receptor_3" title="Somatostatin receptor 3">3</a></li> <li><a href="/wiki/Somatostatin_receptor_4" title="Somatostatin receptor 4">4</a></li> <li><a href="/wiki/Somatostatin_receptor_5" title="Somatostatin receptor 5">5</a>)</li> <li><a href="/wiki/Tachykinin_receptor" title="Tachykinin receptor">Tachykinin</a> (<a href="/wiki/Tachykinin_receptor_1" title="Tachykinin receptor 1">1</a></li> <li><a href="/wiki/Tachykinin_receptor_2" title="Tachykinin receptor 2">2</a></li> <li><a href="/wiki/Tachykinin_receptor_3" title="Tachykinin receptor 3">3</a>)</li> <li><a href="/wiki/Thyrotropin_receptor" title="Thyrotropin receptor">Thyrotropin</a></li> <li><a href="/wiki/Thyrotropin-releasing_hormone_receptor" title="Thyrotropin-releasing hormone receptor">Thyrotropin-releasing hormone</a></li> <li><a href="/wiki/Urotensin-II_receptor" title="Urotensin-II receptor">Urotensin-II</a></li> <li><a href="/wiki/Vasopressin_receptor" title="Vasopressin receptor">Vasopressin</a> (<a href="/wiki/Arginine_vasopressin_receptor_1A" class="mw-redirect" title="Arginine vasopressin receptor 1A">1A</a></li> <li><a href="/wiki/Arginine_vasopressin_receptor_1B" class="mw-redirect" title="Arginine vasopressin receptor 1B">1B</a></li> <li><a href="/wiki/Arginine_vasopressin_receptor_2" class="mw-redirect" title="Arginine vasopressin receptor 2">2</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;">Miscellaneous</th><td class="navbox-list-with-group navbox-list navbox-odd" style="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:7em;text-align:left;"><a href="/wiki/Taste_receptor" title="Taste receptor">Taste</a>, bitter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>TAS2R</i> <ul><li><a href="/wiki/TAS2R1" title="TAS2R1">1</a></li> <li><a href="/wiki/TAS2R3" title="TAS2R3">3</a></li> <li><a href="/wiki/TAS2R4" title="TAS2R4">4</a></li> <li><a href="/wiki/TAS2R5" title="TAS2R5">5</a></li> <li><a href="/wiki/TAS2R7" title="TAS2R7">7</a></li> <li><a href="/wiki/TAS2R8" title="TAS2R8">8</a></li> <li><a href="/wiki/TAS2R9" title="TAS2R9">9</a></li> <li><a href="/wiki/TAS2R10" title="TAS2R10">10</a></li> <li><a href="/wiki/TAS2R13" title="TAS2R13">13</a></li> <li><a href="/wiki/TAS2R14" title="TAS2R14">14</a></li> <li><a href="/wiki/TAS2R16" title="TAS2R16">16</a></li> <li><a href="/wiki/TAS2R19" title="TAS2R19">19</a></li> <li><a href="/wiki/TAS2R20" title="TAS2R20">20</a></li> <li><a href="/wiki/TAS2R30" title="TAS2R30">30</a></li> <li><a href="/wiki/TAS2R31" title="TAS2R31">31</a></li> <li><a href="/wiki/TAS2R38" title="TAS2R38">38</a></li> <li><a href="/wiki/TAS2R39" title="TAS2R39">39</a></li> <li><a href="/wiki/TAS2R40" title="TAS2R40">40</a></li> <li><a href="/wiki/TAS2R41" title="TAS2R41">41</a></li> <li><a href="/wiki/TAS2R42" title="TAS2R42">42</a></li> <li><a href="/wiki/TAS2R43" title="TAS2R43">43</a></li> <li><a href="/wiki/TAS2R45" title="TAS2R45">45</a></li> <li><a href="/wiki/TAS2R46" title="TAS2R46">46</a></li> <li><a href="/wiki/TAS2R50" title="TAS2R50">50</a></li> <li><a href="/wiki/TAS2R60" title="TAS2R60">60</a></li></ul></li> <li><a href="/wiki/Vomeronasal_receptor" title="Vomeronasal receptor">Vomeronasal receptor</a> type 1</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;"><a href="/wiki/Orphan_receptor" title="Orphan receptor">Orphan</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>GPR</i> (<a href="/wiki/GPR1" title="GPR1">1</a></li> <li><a href="/wiki/GPR3" title="GPR3">3</a></li> <li><a href="/wiki/GPR4" title="GPR4">4</a></li> <li><a href="/wiki/GPR6" title="GPR6">6</a></li> <li><a href="/wiki/GPR12" title="GPR12">12</a></li> <li><a href="/wiki/GPR15" title="GPR15">15</a></li> <li><a href="/wiki/GPR17" title="GPR17">17</a></li> <li><a href="/wiki/GPR18" class="mw-redirect" title="GPR18">18</a></li> <li><a href="/wiki/GPR19" title="GPR19">19</a></li> <li><a href="/wiki/GPR20" title="GPR20">20</a></li> <li><a href="/wiki/GPR21" title="GPR21">21</a></li> <li><a href="/wiki/GPR22" title="GPR22">22</a></li> <li><a href="/wiki/LPAR4" title="LPAR4">23</a></li> <li><a href="/wiki/GPR25" title="GPR25">25</a></li> <li><a href="/wiki/GPR26" title="GPR26">26</a></li> <li><a href="/wiki/GPR27" title="GPR27">27</a></li> <li><a href="/wiki/GPR31" title="GPR31">31</a></li> <li><a href="/wiki/GPR32" title="GPR32">32</a></li> <li><a href="/wiki/GPR33" title="GPR33">33</a></li> <li><a href="/wiki/GPR34" title="GPR34">34</a></li> <li><a href="/wiki/GPR35" title="GPR35">35</a></li> <li><a href="/wiki/GPR37" title="GPR37">37</a></li> <li><a href="/wiki/GPR39" title="GPR39">39</a></li> <li><a href="/wiki/GPR42" title="GPR42">42</a></li> <li><a href="/wiki/GPR44" class="mw-redirect" title="GPR44">44</a></li> <li><a href="/wiki/GPR45" title="GPR45">45</a></li> <li><a href="/wiki/GPR50" title="GPR50">50</a></li> <li><a href="/wiki/GPR52" title="GPR52">52</a></li> <li><a href="/wiki/GPR55" title="GPR55">55</a></li> <li><a href="/wiki/GPR61" title="GPR61">61</a></li> <li><a href="/wiki/GPR62" title="GPR62">62</a></li> <li><a href="/wiki/GPR63" title="GPR63">63</a></li> <li><a href="/wiki/GPR65" title="GPR65">65</a></li> <li><a href="/wiki/GPR68" title="GPR68">68</a></li> <li><a href="/wiki/GPR75" title="GPR75">75</a></li> <li><a href="/wiki/GPR78" title="GPR78">78</a></li> <li><a href="/wiki/GPR81" class="mw-redirect" title="GPR81">81</a></li> <li><a href="/wiki/GPR82" title="GPR82">82</a></li> <li><a href="/wiki/GPR83" title="GPR83">83</a></li> <li><a href="/wiki/GPR84" title="GPR84">84</a></li> <li><a href="/wiki/GPR85" title="GPR85">85</a></li> <li><a href="/wiki/GPR87" title="GPR87">87</a></li> <li><a href="/wiki/GPR88" title="GPR88">88</a></li> <li><a href="/wiki/LPAR5" title="LPAR5">92</a></li> <li><a href="/wiki/GPR101" title="GPR101">101</a></li> <li><a href="/wiki/Pyroglutamylated_RFamide_peptide_receptor" title="Pyroglutamylated RFamide peptide receptor">103</a></li> <li><a href="/wiki/Niacin_receptor_1" class="mw-redirect" title="Niacin receptor 1">109A</a></li> <li><a href="/wiki/Niacin_receptor_2" class="mw-redirect" title="Niacin receptor 2">109B</a></li> <li><a href="/wiki/GPR119" title="GPR119">119</a></li> <li><a href="/wiki/GPR120" class="mw-redirect" title="GPR120">120</a></li> <li><a href="/wiki/GPR132" title="GPR132">132</a></li> <li><a href="/wiki/GPR135" title="GPR135">135</a></li> <li><a href="/wiki/GPR137B" title="GPR137B">137B</a></li> <li><a href="/wiki/GPR139" title="GPR139">139</a></li> <li><a href="/wiki/GPR141" title="GPR141">141</a></li> <li><a href="/wiki/GPR142" title="GPR142">142</a></li> <li><a href="/wiki/GPR146" title="GPR146">146</a></li> <li><a href="/wiki/GPR148" title="GPR148">148</a></li> <li><a href="/wiki/GPR149" title="GPR149">149</a></li> <li><a href="/wiki/GPR150" title="GPR150">150</a></li> <li><a href="/wiki/GPR151" title="GPR151">151</a></li> <li><a href="/wiki/GPR152" title="GPR152">152</a></li> <li><a href="/wiki/GPR153" title="GPR153">153</a></li> <li><a href="/wiki/GPR160" title="GPR160">160</a></li> <li><a href="/wiki/GPR161" title="GPR161">161</a></li> <li><a href="/wiki/GPR162" title="GPR162">162</a></li> <li><a href="/wiki/GPR171" title="GPR171">171</a></li> <li><a href="/wiki/GPR173" title="GPR173">173</a></li> <li><a href="/wiki/GPR174" title="GPR174">174</a></li> <li><a href="/wiki/GPR176" title="GPR176">176</a></li> <li><a href="/wiki/GPR177" class="mw-redirect" title="GPR177">177</a></li> <li><a href="/wiki/GPR182" title="GPR182">182</a></li> <li><a href="/wiki/GPR183" title="GPR183">183</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align:left;"><i>Other</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/GPR182" title="GPR182">Adrenomedullin</a></li> <li><a href="/wiki/Olfactory_receptor" title="Olfactory receptor">Olfactory</a></li> <li><a href="/wiki/Opsin" title="Opsin">Opsin</a> (<a href="/wiki/OPN3" title="OPN3">3</a></li> <li><a href="/wiki/Melanopsin" title="Melanopsin">4</a></li> <li><a href="/wiki/OPN5" title="OPN5">5</a></li> <li><a href="/wiki/OPN1LW" title="OPN1LW">1LW</a></li> <li><a href="/wiki/OPN1MW" title="OPN1MW">1MW</a></li> <li><a href="/wiki/OPN1SW" title="OPN1SW">1SW</a></li> <li><a href="/wiki/Retinal_G_protein_coupled_receptor" title="Retinal G protein coupled receptor">RGR</a></li> <li><a href="/wiki/RRH" title="RRH">RRH</a>)</li> <li><a href="/wiki/Protease-activated_receptor" title="Protease-activated receptor">Protease-activated</a> (<a href="/wiki/Coagulation_factor_II_receptor" class="mw-redirect" title="Coagulation factor II receptor">1</a></li> <li><a href="/wiki/Protease_activated_receptor_2" class="mw-redirect" title="Protease activated receptor 2">2</a></li> <li><a href="/wiki/F2RL2" title="F2RL2">3</a></li> <li><a href="/wiki/F2RL3" title="F2RL3">4</a>)</li> <li><a href="/wiki/Super_Conserved_Receptor_Expressed_in_Brain" title="Super Conserved Receptor Expressed in Brain">SREB</a> (<a href="/wiki/GPR27" title="GPR27">1</a></li> <li><a href="/wiki/GPR85" title="GPR85">2</a></li> <li><a href="/wiki/GPR173" title="GPR173">3</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Class_B:_Secretin-like" style="font-size:114%;margin:0 4em"><a href="/wiki/Secretin_receptor_family" title="Secretin receptor family">Class B</a>: <a href="/wiki/Secretin_receptor" title="Secretin receptor">Secretin</a>-like</div></th></tr><tr><td colspan="2" class="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:10em;text-align:left;"><a href="/wiki/Adhesion_G_protein-coupled_receptor" title="Adhesion G protein-coupled receptor">Adhesion</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>ADGRB</i> <ul><li><a href="/wiki/Brain-specific_angiogenesis_inhibitor" title="Brain-specific angiogenesis inhibitor">Brain-specific angiogenesis inhibitor</a> <ul><li><a href="/wiki/Brain-specific_angiogenesis_inhibitor_1" title="Brain-specific angiogenesis inhibitor 1">1</a></li> <li><a href="/wiki/Brain-specific_angiogenesis_inhibitor_2" title="Brain-specific angiogenesis inhibitor 2">2</a></li> <li><a href="/wiki/Brain-specific_angiogenesis_inhibitor_3" title="Brain-specific angiogenesis inhibitor 3">3</a></li></ul></li></ul></li> <li><i>ADGRC</i> <ul><li><a href="/wiki/Flamingo_(protein)" title="Flamingo (protein)">Cadherin</a> <ul><li><a href="/wiki/CELSR1" title="CELSR1">1</a></li> <li><a href="/wiki/CELSR2" title="CELSR2">2</a></li> <li><a href="/wiki/CELSR3" title="CELSR3">3</a></li></ul></li></ul></li> <li><i>ADGRE</i> <ul><li><a href="/wiki/EGF_module-containing_mucin-like_hormone_receptor" title="EGF module-containing mucin-like hormone receptor">EMR</a> <ul><li><a href="/wiki/EMR1" title="EMR1">1</a></li> <li><a href="/wiki/EMR2" title="EMR2">2</a></li> <li><a href="/wiki/EMR3" title="EMR3">3</a></li></ul></li> <li><a href="/wiki/CD97" title="CD97">CD97</a></li></ul></li> <li><i>ADGRG</i> <ul><li><a href="/wiki/GPR56" title="GPR56">1</a></li> <li><a href="/wiki/GPR64" title="GPR64">2</a></li> <li><a href="/wiki/GPR97" title="GPR97">3</a></li> <li><a href="/wiki/GPR112" title="GPR112">4</a></li> <li><a href="/wiki/GPR114" title="GPR114">5</a></li> <li><a href="/wiki/GPR126" title="GPR126">6</a></li> <li><a href="/wiki/GPR128" title="GPR128">7</a></li></ul></li> <li><i>ADGRL</i> <ul><li><a href="/wiki/Latrophilin_receptor" class="mw-redirect" title="Latrophilin receptor">Latrophilin</a> <ul><li><a href="/wiki/Latrophilin_1" title="Latrophilin 1">1</a></li> <li><a href="/wiki/Latrophilin_2" title="Latrophilin 2">2</a></li> <li><a href="/wiki/Latrophilin_3" title="Latrophilin 3">3</a></li></ul></li> <li><a href="/wiki/ELTD1" class="mw-redirect" title="ELTD1">ELTD1</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em;text-align:left;"><a href="/wiki/Orphan_receptor" title="Orphan receptor">Orphan</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>GPR</i> (<a href="/wiki/GPR56" title="GPR56">56</a></li> <li><a href="/wiki/GPR64" title="GPR64">64</a></li> <li><a href="/wiki/GPR97" title="GPR97">97</a></li> <li><a href="/wiki/GPR98" title="GPR98">98</a></li> <li><a href="/wiki/GPR110" title="GPR110">110</a></li> <li><a href="/wiki/GPR111" title="GPR111">111</a></li> <li><a href="/wiki/GPR112" title="GPR112">112</a></li> <li><a href="/wiki/GPR113" title="GPR113">113</a></li> <li><a href="/wiki/GPR114" title="GPR114">114</a></li> <li><a href="/wiki/GPR115" title="GPR115">115</a></li> <li><a href="/wiki/GPR116" title="GPR116">116</a></li> <li><a href="/wiki/GPR123" title="GPR123">123</a></li> <li><a href="/wiki/GPR124" title="GPR124">124</a></li> <li><a href="/wiki/GPR125" title="GPR125">125</a></li> <li><a href="/wiki/GPR126" title="GPR126">126</a></li> <li><a href="/wiki/GPR128" title="GPR128">128</a></li> <li><a href="/wiki/GPR133" title="GPR133">133</a></li> <li><a href="/wiki/GPR143" title="GPR143">143</a></li> <li><a href="/wiki/GPR144" title="GPR144">144</a></li> <li><a href="/wiki/GPR155" title="GPR155">155</a></li> <li><a href="/wiki/GPR157" title="GPR157">157</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em;text-align:left;"><i>Other</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Calcitonin_receptor" title="Calcitonin receptor">Calcitonin</a></li> <li><a href="/wiki/CALCRL" title="CALCRL">CALCRL</a></li> <li><a href="/wiki/Corticotropin-releasing_hormone_receptor" title="Corticotropin-releasing hormone receptor">Corticotropin-releasing hormone</a> (<a href="/wiki/Corticotropin-releasing_hormone_receptor_1" title="Corticotropin-releasing hormone receptor 1">1</a></li> <li><a href="/wiki/Corticotropin-releasing_hormone_receptor_2" title="Corticotropin-releasing hormone receptor 2">2</a>)</li> <li><a href="/wiki/Glucagon_receptor_family" title="Glucagon receptor family">Glucagon</a> (<a href="/wiki/Glucagon_receptor" title="Glucagon receptor">GR</a></li> <li><a href="/wiki/Gastric_inhibitory_polypeptide_receptor" title="Gastric inhibitory polypeptide receptor">GIPR</a></li> <li><a href="/wiki/Glucagon-like_peptide-1_receptor" title="Glucagon-like peptide-1 receptor">GLP1R</a></li> <li><a href="/wiki/Glucagon-like_peptide-2_receptor" title="Glucagon-like peptide-2 receptor">GLP2R</a>)</li> <li><a href="/wiki/Growth-hormone-releasing_hormone_receptor" title="Growth-hormone-releasing hormone receptor">Growth-hormone-releasing hormone</a></li> <li><a href="/wiki/ADCYAP1R1" title="ADCYAP1R1">PACAPR1</a></li> <li><a href="/wiki/Orphan_receptor" title="Orphan receptor">GPR</a></li> <li><a href="/wiki/Methuselah-like_proteins" title="Methuselah-like proteins">Methuselah-like proteins</a></li> <li><a href="/wiki/Parathyroid_hormone_receptor" title="Parathyroid hormone receptor">Parathyroid hormone</a> (<a href="/wiki/Parathyroid_hormone_1_receptor" title="Parathyroid hormone 1 receptor">1</a></li> <li><a href="/wiki/Parathyroid_hormone_2_receptor" title="Parathyroid hormone 2 receptor">2</a>)</li> <li><a href="/wiki/Secretin_receptor" title="Secretin receptor">Secretin</a></li> <li><a href="/wiki/Vasoactive_intestinal_peptide_receptor" title="Vasoactive intestinal peptide receptor">Vasoactive intestinal peptide</a> (<a href="/wiki/VIPR1" title="VIPR1">1</a></li> <li><a href="/wiki/VIPR2" title="VIPR2">2</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Class_C:_Metabotropic_glutamate_/_pheromone" style="font-size:114%;margin:0 4em"><a href="/wiki/Class_C_GPCR" title="Class C GPCR">Class C</a>: <a href="/wiki/Metabotropic_receptor" title="Metabotropic receptor">Metabotropic</a> <a href="/wiki/Metabotropic_glutamate_receptor" title="Metabotropic glutamate receptor">glutamate</a> / <a href="/wiki/Pheromone" title="Pheromone">pheromone</a></div></th></tr><tr><td colspan="2" class="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:10em;text-align:left;"><a href="/wiki/Taste_receptor" title="Taste receptor">Taste</a>, sweet</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>TAS1R</i> <ul><li><a href="/wiki/TAS1R1" title="TAS1R1">1</a></li> <li><a href="/wiki/TAS1R2" title="TAS1R2">2</a></li> <li><a href="/wiki/TAS1R3" title="TAS1R3">3</a></li></ul></li> <li><a href="/wiki/Vomeronasal_receptor" title="Vomeronasal receptor">Vomeronasal receptor</a>, type 2</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em;text-align:left;"><i>Other</i></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Calcium-sensing_receptor" title="Calcium-sensing receptor">Calcium-sensing receptor</a></li> <li><a href="/wiki/GABA_receptor" title="GABA receptor">GABA<sub>B</sub></a> (<a href="/wiki/GABBR1" title="GABBR1">1</a></li> <li><a href="/wiki/GABBR2" title="GABBR2">2</a>)</li> <li><a href="/wiki/Glutamate_receptor" title="Glutamate receptor">Glutamate receptor</a> (<a href="/wiki/Metabotropic_glutamate_receptor" title="Metabotropic glutamate receptor">Metabotropic glutamate</a> (<a href="/wiki/Metabotropic_glutamate_receptor_1" title="Metabotropic glutamate receptor 1">1</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_2" title="Metabotropic glutamate receptor 2">2</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_3" title="Metabotropic glutamate receptor 3">3</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_4" title="Metabotropic glutamate receptor 4">4</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_5" title="Metabotropic glutamate receptor 5">5</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_6" title="Metabotropic glutamate receptor 6">6</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_7" title="Metabotropic glutamate receptor 7">7</a></li> <li><a href="/wiki/Metabotropic_glutamate_receptor_8" title="Metabotropic glutamate receptor 8">8</a>))</li> <li><a href="/wiki/GPRC6A" title="GPRC6A">GPRC6A</a></li> <li><a href="/wiki/Orphan_receptor" title="Orphan receptor">GPR</a> (<a href="/wiki/GPR156" title="GPR156">156</a></li> <li><a href="/wiki/GPR158" title="GPR158">158</a></li> <li><a href="/wiki/GPR179" title="GPR179">179</a>)</li> <li><a href="/wiki/Retinoic_acid-inducible_orphan_G_protein-coupled_receptor" title="Retinoic acid-inducible orphan G protein-coupled receptor">RAIG</a> (<a href="/wiki/GPRC5A" title="GPRC5A">1</a></li> <li><a href="/wiki/GPRC5B" title="GPRC5B">2</a></li> <li><a href="/wiki/GPRC5C" title="GPRC5C">3</a></li> <li><a href="/wiki/GPRC5D" title="GPRC5D">4</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Class_F:_Frizzled_&amp;_Smoothened" style="font-size:114%;margin:0 4em">Class F: <a href="/wiki/Frizzled" title="Frizzled">Frizzled</a> & <a href="/wiki/Smoothened" title="Smoothened">Smoothened</a></div></th></tr><tr><td colspan="2" class="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:10em;text-align:left;">Frizzled</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Frizzled" title="Frizzled">Frizzled</a> (<a href="/wiki/FZD1" class="mw-redirect" title="FZD1">1</a></li> <li><a href="/wiki/FZD2" class="mw-redirect" title="FZD2">2</a></li> <li><a href="/wiki/FZD3" class="mw-redirect" title="FZD3">3</a></li> <li><a href="/wiki/FZD4" class="mw-redirect" title="FZD4">4</a></li> <li><a href="/wiki/FZD5" class="mw-redirect" title="FZD5">5</a></li> <li><a href="/wiki/FZD6" class="mw-redirect" title="FZD6">6</a></li> <li><a href="/wiki/FZD7" class="mw-redirect" title="FZD7">7</a></li> <li><a href="/wiki/FZD8" class="mw-redirect" title="FZD8">8</a></li> <li><a href="/wiki/FZD9" class="mw-redirect" title="FZD9">9</a></li> <li><a href="/wiki/FZD10" class="mw-redirect" title="FZD10">10</a>)</li></ul> </div></td></tr><tr><th scope="row" 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