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vector-toc-level-2"> <a class="vector-toc-link" href="#GluN2"> <div class="vector-toc-text"> 5.2 GluN2 </div> </a> <ul id="toc-GluN2-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-GluN2B_to_GluN2A_switch" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#GluN2B_to_GluN2A_switch"> <div class="vector-toc-text"> 5.3 GluN2B to GluN2A switch </div> </a> <ul id="toc-GluN2B_to_GluN2A_switch-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-GluN2B_to_GluN2C_switch" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#GluN2B_to_GluN2C_switch"> <div class="vector-toc-text"> 5.4 GluN2B to GluN2C switch </div> </a> <ul id="toc-GluN2B_to_GluN2C_switch-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Role_in_excitotoxicity" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_excitotoxicity"> <div class="vector-toc-text"> 6 Role in excitotoxicity </div> </a> <button aria-controls="toc-Role_in_excitotoxicity-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Role in excitotoxicity subsection </button> <ul id="toc-Role_in_excitotoxicity-sublist" class="vector-toc-list"> <li id="toc-Differing_cascade_pathways" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Differing_cascade_pathways"> <div class="vector-toc-text"> 6.1 Differing cascade pathways </div> </a> <ul id="toc-Differing_cascade_pathways-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neural_plasticity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neural_plasticity"> <div class="vector-toc-text"> 6.2 Neural plasticity </div> </a> <ul id="toc-Neural_plasticity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_of_differing_subunits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_of_differing_subunits"> <div class="vector-toc-text"> 6.3 Role of differing subunits </div> </a> <ul id="toc-Role_of_differing_subunits-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Excitotoxicity_in_a_clinical_setting" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Excitotoxicity_in_a_clinical_setting"> <div class="vector-toc-text"> 6.4 Excitotoxicity in a clinical setting </div> </a> <ul id="toc-Excitotoxicity_in_a_clinical_setting-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Ligands" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Ligands"> <div class="vector-toc-text"> 7 Ligands </div> </a> <button aria-controls="toc-Ligands-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Ligands subsection </button> <ul id="toc-Ligands-sublist" class="vector-toc-list"> <li id="toc-Agonists" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Agonists"> <div class="vector-toc-text"> 7.1 Agonists </div> </a> <ul id="toc-Agonists-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"> 7.1.1 Examples </div> </a> <ul id="toc-Examples-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neramexane" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Neramexane"> <div class="vector-toc-text"> 7.1.2 Neramexane </div> </a> <ul id="toc-Neramexane-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Partial_agonists" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Partial_agonists"> <div class="vector-toc-text"> 7.2 Partial agonists </div> </a> <ul id="toc-Partial_agonists-sublist" class="vector-toc-list"> <li id="toc-Examples_2" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Examples_2"> <div class="vector-toc-text"> 7.2.1 Examples </div> </a> <ul id="toc-Examples_2-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Antagonists" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Antagonists"> <div class="vector-toc-text"> 7.3 Antagonists </div> </a> <ul id="toc-Antagonists-sublist" class="vector-toc-list"> <li id="toc-Examples_3" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Examples_3"> <div class="vector-toc-text"> 7.3.1 Examples </div> </a> <ul id="toc-Examples_3-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nitromemantine" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Nitromemantine"> <div class="vector-toc-text"> 7.3.2 Nitromemantine </div> </a> <ul id="toc-Nitromemantine-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Modulators" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Modulators"> <div class="vector-toc-text"> 7.4 Modulators </div> </a> <ul id="toc-Modulators-sublist" class="vector-toc-list"> <li id="toc-Examples_4" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Examples_4"> <div class="vector-toc-text"> 7.4.1 Examples </div> </a> <ul id="toc-Examples_4-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Development_of_NMDA_receptor_antagonists" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Development_of_NMDA_receptor_antagonists"> <div class="vector-toc-text"> 7.5 Development of NMDA receptor antagonists </div> </a> <ul id="toc-Development_of_NMDA_receptor_antagonists-sublist" class="vector-toc-list"> <li id="toc-Competitive_NMDA_receptor_antagonists" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Competitive_NMDA_receptor_antagonists"> <div class="vector-toc-text"> 7.5.1 Competitive NMDA receptor antagonists </div> </a> <ul id="toc-Competitive_NMDA_receptor_antagonists-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Noncompetitive_NMDA_receptor_antagonists" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Noncompetitive_NMDA_receptor_antagonists"> <div class="vector-toc-text"> 7.5.2 Noncompetitive NMDA receptor antagonists </div> </a> <ul id="toc-Noncompetitive_NMDA_receptor_antagonists-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Memantine_and_related_compounds" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Memantine_and_related_compounds"> <div class="vector-toc-text"> 7.5.3 Memantine and related compounds </div> </a> <ul id="toc-Memantine_and_related_compounds-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Structure_activity_relationship_(SAR)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structure_activity_relationship_(SAR)"> <div class="vector-toc-text"> 7.6 Structure activity relationship (SAR) </div> </a> <ul id="toc-Structure_activity_relationship_(SAR)-sublist" class="vector-toc-list"> <li id="toc-Second_generation_derivative_of_memantine;_nitromemantine" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Second_generation_derivative_of_memantine;_nitromemantine"> <div class="vector-toc-text"> 7.6.1 Second generation derivative of memantine; nitromemantine </div> </a> <ul id="toc-Second_generation_derivative_of_memantine;_nitromemantine-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Therapeutic_application" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Therapeutic_application"> <div class="vector-toc-text"> 7.7 Therapeutic application </div> </a> <ul id="toc-Therapeutic_application-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Receptor_modulation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Receptor_modulation"> <div class="vector-toc-text"> 8 Receptor modulation </div> </a> <ul id="toc-Receptor_modulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> 9 Clinical significance </div> </a> <ul id="toc-Clinical_significance-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"> 10 See also </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"> 11 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 12 External links </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 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Available in 22 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-22" aria-hidden="true" > 22 languages </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%86-%D9%85%D8%AB%D9%8A%D9%84-%D8%AF-%D8%A3%D8%B3%D8%A8%D8%A7%D8%B1%D8%AA%D8%A7%D8%AA" title="مستقبل ن-مثيل-د-أسبارتات – Arabic" lang="ar" hreflang="ar" data-title="مستقبل ن-مثيل-د-أسبارتات" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Receptor_NMDA" title="Receptor NMDA – Catalan" lang="ca" hreflang="ca" data-title="Receptor NMDA" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/NMDA_receptor" title="NMDA receptor – Czech" lang="cs" hreflang="cs" data-title="NMDA receptor" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/NMDA-Rezeptor" title="NMDA-Rezeptor – German" lang="de" hreflang="de" data-title="NMDA-Rezeptor" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Receptor_NMDA" title="Receptor NMDA – Spanish" lang="es" hreflang="es" data-title="Receptor NMDA" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</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_NMDA" title="گیرنده NMDA – Persian" lang="fa" hreflang="fa" data-title="گیرنده NMDA" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/R%C3%A9cepteur_NMDA" title="Récepteur NMDA – French" lang="fr" hreflang="fr" data-title="Récepteur NMDA" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/NMDA_%EC%88%98%EC%9A%A9%EC%B2%B4" title="NMDA 수용체 – Korean" lang="ko" hreflang="ko" data-title="NMDA 수용체" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Recettore_NMDA" title="Recettore NMDA – Italian" lang="it" hreflang="it" data-title="Recettore NMDA" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Receptorium_NMDA" title="Receptorium NMDA – Latin" lang="la" hreflang="la" data-title="Receptorium NMDA" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/NMDA-receptor" title="NMDA-receptor – Dutch" lang="nl" hreflang="nl" data-title="NMDA-receptor" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/NMDA%E5%9E%8B%E3%82%B0%E3%83%AB%E3%82%BF%E3%83%9F%E3%83%B3%E9%85%B8%E5%8F%97%E5%AE%B9%E4%BD%93" title="NMDA型グルタミン酸受容体 – Japanese" lang="ja" hreflang="ja" data-title="NMDA型グルタミン酸受容体" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Receptor_NMDA" title="Receptor NMDA – Polish" lang="pl" hreflang="pl" data-title="Receptor NMDA" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Receptor_NMDA" title="Receptor NMDA – Portuguese" lang="pt" hreflang="pt" data-title="Receptor NMDA" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/NMDA-%D1%80%D0%B5%D1%86%D0%B5%D0%BF%D1%82%D0%BE%D1%80" title="NMDA-рецептор – Russian" lang="ru" hreflang="ru" data-title="NMDA-рецептор" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/NMDA_receptor" title="NMDA receptor – Serbian" lang="sr" hreflang="sr" data-title="NMDA receptor" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/NMDA_receptor" title="NMDA receptor – Serbo-Croatian" lang="sh" hreflang="sh" data-title="NMDA receptor" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/NMDA-reseptori" title="NMDA-reseptori – Finnish" lang="fi" hreflang="fi" data-title="NMDA-reseptori" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/NMDA-receptorn" title="NMDA-receptorn – Swedish" lang="sv" hreflang="sv" data-title="NMDA-receptorn" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/NMDA_resept%C3%B6r%C3%BC" title="NMDA reseptörü – Turkish" lang="tr" hreflang="tr" data-title="NMDA reseptörü" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/NMDA-%D1%80%D0%B5%D1%86%D0%B5%D0%BF%D1%82%D0%BE%D1%80" title="NMDA-рецептор – Ukrainian" lang="uk" hreflang="uk" data-title="NMDA-рецептор" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/N-%E7%94%B2%E5%9F%BA-D-%E5%A4%A9%E9%96%80%E5%86%AC%E8%83%BA%E9%85%B8%E5%8F%97%E9%AB%94" title="N-甲基-D-天門冬胺酸受體 – Chinese" lang="zh" hreflang="zh" data-title="N-甲基-D-天門冬胺酸受體" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li> </ul> <div class="after-portlet after-portlet-lang"><a 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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">"NR1" redirects here. For the submarine, see <a href="/wiki/NR-1" class="mw-redirect" title="NR-1">NR-1</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/NDMA_(disambiguation)" class="mw-redirect mw-disambig" title="NDMA (disambiguation)">NDMA (disambiguation)</a>.</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Activated_NMDAR.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Activated_NMDAR.svg/220px-Activated_NMDAR.svg.png" decoding="async" width="220" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Activated_NMDAR.svg/330px-Activated_NMDAR.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/00/Activated_NMDAR.svg/440px-Activated_NMDAR.svg.png 2x" data-file-width="303" data-file-height="312" /></a><figcaption>Stylized depiction of an activated NMDAR. Glutamate is in the glutamate-binding site and glycine is in the glycine-binding site. The <a href="/wiki/Allosteric_site" class="mw-redirect" title="Allosteric site">allosteric site</a>, which modulates receptor function when bound to a ligand, is not occupied. NMDARs require the binding of two molecules of <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> or <a href="/wiki/Aspartate" class="mw-redirect" title="Aspartate">aspartate</a> and two of <a href="/wiki/Glycine" title="Glycine">glycine</a><a href="#cite_note-Laube-1">[1]</a><a href="#cite_note-Anson-2">[2]</a></figcaption></figure> The N-methyl-D-aspartate receptor (also known as the NMDA receptor or NMDAR), is a <a href="/wiki/Glutamate_receptor" title="Glutamate receptor">glutamate receptor</a> and predominantly Ca2+ <a href="/wiki/Ion_channel" title="Ion channel">ion channel</a> found in <a href="/wiki/Neuron" title="Neuron">neurons</a>.<a href="#cite_note-3">[3]</a><a href="#cite_note-4">[4]</a> The NMDA receptor is one of three types of <a href="/wiki/Ionotropic_glutamate_receptor" title="Ionotropic glutamate receptor">ionotropic glutamate receptors</a>, the other two being <a href="/wiki/AMPA_receptor" title="AMPA receptor">AMPA</a> and <a href="/wiki/Kainate_receptor" title="Kainate receptor">kainate receptors</a>. Depending on its subunit composition, its <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligands</a> are <a href="/wiki/Glutamate_(neurotransmitter)" title="Glutamate (neurotransmitter)">glutamate</a> and <a href="/wiki/Glycine" title="Glycine">glycine</a> (or <a href="/wiki/D-Serine" class="mw-redirect" title="D-Serine">D-serine</a>). However, the binding of the ligands is typically not sufficient to open the channel as it may be blocked by <a href="/wiki/Magnesium" title="Magnesium">Mg2+</a> ions which are only removed when the neuron is sufficiently depolarized. Thus, the channel acts as a "coincidence detector" and only once both of these conditions are met, the channel opens and it allows <a href="/wiki/Cation" class="mw-redirect" title="Cation">positively charged ions</a> (cations) to flow through the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a>.<a href="#cite_note-Furukawa-5">[5]</a> The NMDA receptor is thought to be very important for controlling <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a> and mediating <a href="/wiki/Learning" title="Learning">learning</a> and <a href="/wiki/Memory" title="Memory">memory</a> functions.<a href="#cite_note-pmid19605837-6">[6]</a> The NMDA receptor is <a href="/wiki/Ionotropic" class="mw-redirect" title="Ionotropic">ionotropic</a>, meaning it is a protein which allows the passage of ions through the cell membrane.<a href="#cite_note-pmid1834949-7">[7]</a> The NMDA receptor is so named because the <a href="/wiki/Agonist" title="Agonist">agonist</a> molecule <a href="/wiki/N-methyl-D-aspartate" class="mw-redirect" title="N-methyl-D-aspartate">N-methyl-D-aspartate</a> (NMDA) binds selectively to it, and not to other <a href="/wiki/Glutamate_receptor" title="Glutamate receptor">glutamate receptors</a>. Activation of NMDA receptors results in the opening of the ion channel that is nonselective to <a href="/wiki/Ion" title="Ion">cations</a>, with a combined <a href="/wiki/Reversal_potential" title="Reversal potential">reversal potential</a> near 0 mV. While the opening and closing of the ion channel is primarily gated by ligand binding, the current flow through the ion channel is voltage-dependent. Specifically located on the receptor, extracellular magnesium (Mg2+) and zinc (Zn2+) ions can bind and prevent other cations from flowing through the open ion channel. A voltage-dependent flow of predominantly calcium (Ca2+), sodium (Na+), and potassium (K+) ions into and out of the cell is made possible by the depolarization of the cell, which displaces and repels the Mg2+ and Zn2+ ions from the pore.<a href="#cite_note-pmid10049997-8">[8]</a><a href="#cite_note-pmid11775847-9">[9]</a><a href="#cite_note-pmid11399431-10">[10]</a><a href="#cite_note-pmid17088105-11">[11]</a> Ca2+ flux through NMDA receptors in particular is thought to be critical in synaptic plasticity, a cellular mechanism for learning and memory, due to proteins which bind to and are activated by Ca2+ ions. Activity of the NMDA receptor is blocked by many <a href="/wiki/Psychoactive" class="mw-redirect" title="Psychoactive">psychoactive</a> drugs such as <a href="/wiki/Phencyclidine" title="Phencyclidine">phencyclidine</a> (PCP), <a href="/wiki/Alcohol_(drug)" title="Alcohol (drug)">alcohol</a> (<a href="/wiki/Ethanol" title="Ethanol">ethanol</a>) and <a href="/wiki/Dextromethorphan" title="Dextromethorphan">dextromethorphan</a> (DXM). The <a href="/wiki/Anaesthetic" class="mw-redirect" title="Anaesthetic">anaesthetic</a> and <a href="/wiki/Analgesic" title="Analgesic">analgesic</a> effects of the drugs <a href="/wiki/Ketamine" title="Ketamine">ketamine</a> and <a href="/wiki/Nitrous_oxide" title="Nitrous oxide">nitrous oxide</a> are also partially due to their effects at blocking NMDA receptor activity. In contrast, overactivation of NMDAR by NMDA agonists increases the <a href="/wiki/Cytosol" title="Cytosol">cytosolic</a> concentrations of <a href="/wiki/Calcium" title="Calcium">calcium</a> and <a href="/wiki/Zinc" title="Zinc">zinc</a>, which significantly contributes to <a href="/wiki/Neuron" title="Neuron">neural</a> <a href="/wiki/Neurodegeneration" class="mw-redirect" title="Neurodegeneration">death</a>, an effect known to be prevented by <a href="/wiki/Cannabinoid" title="Cannabinoid">cannabinoids</a>, mediated by activation of the <a href="/wiki/Cannabinoid_receptor_type_1" class="mw-redirect" title="Cannabinoid receptor type 1">CB1 receptor</a>, which leads <a href="/wiki/HINT1" title="HINT1">HINT1</a> protein to counteract the toxic effects of NMDAR-mediated <a href="/wiki/Nitric_oxide" title="Nitric oxide">NO</a> production and zinc release.<a href="#cite_note-12">[12]</a> As well as preventing <a href="/wiki/Methamphetamine" title="Methamphetamine">methamphetamine</a>-induced <a href="/wiki/Neurotoxicity" title="Neurotoxicity">neurotoxicity</a> via inhibition of <a href="/wiki/Nitric_oxide_synthase" title="Nitric oxide synthase">nitric oxide synthase</a> (nNOS) expression and <a href="/wiki/Astrocyte" title="Astrocyte">astrocyte</a> activation, it is seen to reduce methamphetamine induced brain damage through CB1-dependent and independent mechanisms, respectively, and inhibition of methamphetamine induced <a href="/wiki/Astrogliosis" title="Astrogliosis">astrogliosis</a> is likely to occur through a <a href="/wiki/Cannabinoid_receptor_type_2" class="mw-redirect" title="Cannabinoid receptor type 2">CB2 receptor</a> dependent mechanism for <a href="/wiki/Tetrahydrocannabinol" title="Tetrahydrocannabinol">THC</a>.<a href="#cite_note-13">[13]</a> Since 1989, <a href="/wiki/Memantine" title="Memantine">memantine</a> has been recognized to be an <a href="/wiki/Uncompetitive_antagonist" class="mw-redirect" title="Uncompetitive antagonist">uncompetitive antagonist</a> of the NMDA receptor, entering the channel of the receptor after it has been activated and thereby blocking the flow of ions.<a href="#cite_note-Johnson-14">[14]</a><a href="#cite_note-Dominguez-15">[15]</a><a href="#cite_note-Chen-16">[16]</a> Overactivation of the receptor, causing excessive influx of Ca2+ can lead to <a href="/wiki/Excitotoxicity" title="Excitotoxicity">excitotoxicity</a> which is implied to be involved in some neurodegenerative disorders. Blocking of NMDA receptors could therefore, in theory, be useful in treating such diseases.<a href="#cite_note-Chen-16">[16]</a><a href="#cite_note-Kemp-17">[17]</a><a href="#cite_note-Lipton1-18">[18]</a><a href="#cite_note-Koch-19">[19]</a> However, hypofunction of NMDA receptors (due to <a href="/wiki/Glutathione" title="Glutathione">glutathione</a> deficiency or other causes) may be involved in impairment of synaptic plasticity<a href="#cite_note-pmid16330153-20">[20]</a> and could have other negative repercussions. The main problem with the utilization of <a href="/wiki/NMDA_receptor_antagonist" title="NMDA receptor antagonist">NMDA receptor antagonists</a> for <a href="/wiki/Neuroprotection" title="Neuroprotection">neuroprotection</a> is that the physiological actions of the NMDA receptor are essential for normal neuronal function. To be clinically useful NMDA antagonists need to block excessive activation without interfering with normal functions. <a href="/wiki/Memantine" title="Memantine">Memantine</a> has this property.<a href="#cite_note-Lipton2-21">[21]</a> <style data-mw-deduplicate="TemplateStyles:r886046785">.mw-parser-output .toclimit-2 .toclevel-1 ul,.mw-parser-output .toclimit-3 .toclevel-2 ul,.mw-parser-output .toclimit-4 .toclevel-3 ul,.mw-parser-output .toclimit-5 .toclevel-4 ul,.mw-parser-output .toclimit-6 .toclevel-5 ul,.mw-parser-output .toclimit-7 .toclevel-6 ul{display:none}</style><div class="toclimit-3"><meta property="mw:PageProp/toc" /></div> <div class="mw-heading mw-heading2"><h2 id="History">History</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=1" title="Edit section: History">edit</a>]</div> The discovery of NMDA receptors was followed by the synthesis and study of N-methyl-D-aspartic acid (NMDA) in the 1960s by <a href="/w/index.php?title=Jeff_Watkins&action=edit&redlink=1" class="new" title="Jeff Watkins (page does not exist)">Jeff Watkins</a> and colleagues. In the early 1980s, NMDA receptors were shown to be involved in several central synaptic pathways.<a href="#cite_note-Cheng-22">[22]</a><a href="#cite_note-Watkins-23">[23]</a> Receptor subunit selectivity was discovered in the early 1990s, which led to recognition of a new class of compounds that selectively inhibit the <a href="/wiki/NR2B" class="mw-redirect" title="NR2B">NR2B</a> subunit. These findings led to vigorous campaign in the pharmaceutical industry.<a href="#cite_note-pmid17088105-11">[11]</a> From this it was considered that NMDA receptors were associated with a variety of <a href="/wiki/Neurological_disorders" class="mw-redirect" title="Neurological disorders">neurological disorders</a> such as <a href="/wiki/Epilepsy" title="Epilepsy">epilepsy</a>, <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's</a>, <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's</a>, <a href="/wiki/Huntington%27s_disease" title="Huntington's disease">Huntington's</a> and other CNS disorders.<a href="#cite_note-pmid10049997-8">[8]</a> In 2002, it was discovered by <a href="/wiki/Hilmar_Bading" title="Hilmar Bading">Hilmar Bading</a> and co-workers that the cellular consequences of NMDA receptor stimulation depend on the receptor's location on the neuronal cell surface.<a href="#cite_note-pmid11953750-24">[24]</a><a href="#cite_note-:4-25">[25]</a> Synaptic NMDA receptors promote gene expression, plasticity-related events, and acquired <a href="/wiki/Neuroprotection" title="Neuroprotection">neuroprotection</a>. Extrasynaptic NMDA receptors promote death signaling; they cause transcriptional shut-off, mitochondrial dysfunction, and structural disintegration.<a href="#cite_note-pmid11953750-24">[24]</a><a href="#cite_note-:4-25">[25]</a> This pathological triad of extrasynaptic NMDA receptor signaling represents a common conversion point in the etiology of several acute and chronic neurodegenerative conditions.<a href="#cite_note-26">[26]</a> The molecular basis for toxic extrasynaptic NMDA receptor signaling was uncovered by Hilmar Bading and co-workers in 2020.<a href="#cite_note-:5-27">[27]</a> <a href="/wiki/Extrasynaptic_NMDA_receptors" class="mw-redirect" title="Extrasynaptic NMDA receptors">Extrasynaptic NMDA receptors</a> form a death signaling complex with TRPM4. <a href="/w/index.php?title=NMDAR/TRPM4_interaction_interface_inhibitors&action=edit&redlink=1" class="new" title="NMDAR/TRPM4 interaction interface inhibitors (page does not exist)">NMDAR/TRPM4 interaction interface inhibitors</a> (also known as interface inhibitors) disrupt the NMDAR/TRPM4 complex and detoxify extrasynaptic NMDA receptors.<a href="#cite_note-:5-27">[27]</a> A fortuitous finding was made in 1968 when a woman was taking <a href="/wiki/Amantadine" title="Amantadine">amantadine</a> as flu medicine and experienced remarkable remission of her Parkinson's symptoms. This finding, reported by Scawab et al., was the beginning of <a href="/wiki/Medicinal_chemistry" title="Medicinal chemistry">medicinal chemistry</a> of adamantane derivatives in the context of diseases affecting the CNS.<a href="#cite_note-Wanka-28">[28]</a> Before this finding, memantine, another adamantane derivative, had been synthesized by Eli Lilly and Company in 1963. The purpose was to develop a <a href="/wiki/Hypoglycemic" class="mw-redirect" title="Hypoglycemic">hypoglycemic</a> drug, but it showed no such <a href="/wiki/Efficacy" title="Efficacy">efficacy</a>. It was not until 1972 that a possible therapeutic importance of memantine for treating neurodegenerative disorders was discovered. From 1989 memantine has been recognized to be an uncompetitive antagonist of the NMDA receptor.<a href="#cite_note-Dominguez-15">[15]</a> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=2" title="Edit section: Structure">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:7eu7_NMDA-Rezeptor_Regenbogen.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/7eu7_NMDA-Rezeptor_Regenbogen.png/220px-7eu7_NMDA-Rezeptor_Regenbogen.png" decoding="async" width="220" height="274" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/7eu7_NMDA-Rezeptor_Regenbogen.png/330px-7eu7_NMDA-Rezeptor_Regenbogen.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/85/7eu7_NMDA-Rezeptor_Regenbogen.png/440px-7eu7_NMDA-Rezeptor_Regenbogen.png 2x" data-file-width="551" data-file-height="686" /></a><figcaption>Cartoon representation of the human NMDA receptor. Each subunit is individually rainbow colored.</figcaption></figure> Functional NMDA receptors are heterotetramers comprising different combinations of the GluN1, GluN2 (A-D), and GluN3 (A-B) subunits derived from distinct gene families (Grin1-Grin3). All NMDARs contain two of the obligatory GluN1 subunits, which when assembled with GluN2 subunits of the same type, give rise to canonical diheteromeric (d-) NMDARs (e.g., GluN1-2A-1-2A). Triheteromeric NMDARs, by contrast, contain three different types of subunits (e.g., GluN1-2A-1-2B), and include receptors that are composed of one or more subunits from each of the three gene families, designated t-NMDARs (e.g., GluN1-2A-3A-2A).<a href="#cite_note-29">[29]</a> There is one GluN1, four GluN2, and two GluN3 subunit encoding genes, and each gene may produce more than one splice variant. <ul><li>GluN1 – <a href="/wiki/GRIN1" title="GRIN1">GRIN1</a></li> <li>GluN2 <ul><li>GluN2A – <a href="/wiki/GRIN2A" title="GRIN2A">GRIN2A</a></li> <li>GluN2B – <a href="/wiki/GRIN2B" title="GRIN2B">GRIN2B</a></li> <li>GluN2C – <a href="/wiki/GRIN2C" title="GRIN2C">GRIN2C</a></li> <li>GluN2D – <a href="/wiki/GRIN2D" title="GRIN2D">GRIN2D</a></li></ul></li> <li>GluN3 <ul><li>GluN3A – <a href="/wiki/GRIN3A" title="GRIN3A">GRIN3A</a></li> <li>GluN3B – <a href="/wiki/GRIN3B" title="GRIN3B">GRIN3B</a></li></ul></li></ul> <div class="mw-heading mw-heading2"><h2 id="Gating">Gating</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=3" title="Edit section: Gating">edit</a>]</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:N1_N2_NMDA_receptor.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e7/N1_N2_NMDA_receptor.svg/400px-N1_N2_NMDA_receptor.svg.png" decoding="async" width="400" height="400" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e7/N1_N2_NMDA_receptor.svg/600px-N1_N2_NMDA_receptor.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e7/N1_N2_NMDA_receptor.svg/800px-N1_N2_NMDA_receptor.svg.png 2x" data-file-width="500" data-file-height="500" /></a><figcaption>Figure 1: NR1/NR2 NMDA receptor</figcaption></figure> The NMDA receptor is a <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamate</a> and <a href="/wiki/Ion_channel" title="Ion channel">ion channel</a> protein receptor that is activated when <a href="/wiki/Glycine" title="Glycine">glycine</a> and glutamate bind to it.<a href="#cite_note-Furukawa-5">[5]</a> The receptor is a highly complex and dynamic heteromeric protein that interacts with a multitude of intracellular <a href="/wiki/Protein" title="Protein">proteins</a> via three distinct subunits, namely GluN1, GluN2, and GluN3. The GluN1 subunit, which is encoded by the GRIN1 gene, exhibits eight distinct isoforms owing to alternative splicing. On the other hand, the GluN2 subunit, of which there are four different types (A-D), as well as the GluN3 subunit, of which there are two types (A and B), are each encoded by six separate genes. This intricate molecular structure and genetic diversity enable the receptor to carry out a wide range of physiological functions within the <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a>.<a href="#cite_note-Loftis-30">[30]</a><a href="#cite_note-Kristiansen-31">[31]</a> All the subunits share a common membrane topology that is dominated by a large extracellular N-terminus, a membrane region comprising three transmembrane segments, a re-entrant pore loop, an extracellular loop between the transmembrane segments that are structurally not well known, and an intracellular C-terminus, which are different in size depending on the subunit and provide multiple sites of interaction with many intracellular proteins.<a href="#cite_note-Loftis-30">[30]</a><a href="#cite_note-Limapichat-32">[32]</a> Figure 1 shows a basic structure of GluN1/GluN2 subunits that forms the <a href="/wiki/Binding_site" title="Binding site">binding site</a> for memantine, Mg2+ and <a href="/wiki/Ketamine" title="Ketamine">ketamine</a>. <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:NR1-NR2B_subunit.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/44/NR1-NR2B_subunit.png/270px-NR1-NR2B_subunit.png" decoding="async" width="270" height="192" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/44/NR1-NR2B_subunit.png/405px-NR1-NR2B_subunit.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/44/NR1-NR2B_subunit.png/540px-NR1-NR2B_subunit.png 2x" data-file-width="572" data-file-height="406" /></a><figcaption>Figure 2: Transmembrane region of NR1 (left) and NR2B (right) subunits of NMDA receptor</figcaption></figure> Mg2+ blocks the NMDA receptor channel in a voltage-dependent manner. The channels are also highly permeable to Ca2+. Activation of the receptor depends on glutamate binding, <a href="/wiki/D-Serine" class="mw-redirect" title="D-Serine">D-serine</a> or glycine binding at its GluN1-linked binding site and <a href="/wiki/AMPA_receptor" title="AMPA receptor">AMPA receptor</a>-mediated <a href="/wiki/Depolarization" title="Depolarization">depolarization</a> of the postsynaptic membrane, which relieves the voltage-dependent channel block by Mg2+. Activation and opening of the receptor channel thus allows the flow of K+, Na+ and Ca2+ ions, and the influx of Ca2+ triggers intracellular signaling pathways.<a href="#cite_note-Johnson-14">[14]</a><a href="#cite_note-Maher-33">[33]</a> Allosteric receptor binding sites for zinc, proteins and the polyamines spermidine and spermine are also modulators for the NMDA receptor channels.<a href="#cite_note-Danysz-34">[34]</a> The GluN2B subunit has been involved in modulating activity such as learning, memory, processing and feeding behaviors, as well as being implicated in number of human derangements. The basic structure and functions associated with the NMDA receptor can be attributed to the GluN2B subunit. For example, the glutamate binding site and the control of the Mg2+ block are formed by the GluN2B subunit. The high affinity sites for glycine <a href="/wiki/Antagonist" title="Antagonist">antagonist</a> are also exclusively displayed by the GluN1/GluN2B receptor.<a href="#cite_note-Kristiansen-31">[31]</a> GluN1/GluN2B transmembrane segments are considered to be the part of the receptor that forms the binding pockets for uncompetitive NMDA receptor antagonists, but the transmembrane segments structures are not fully known as stated above. It is claimed that three binding sites within the receptor, A644 on the GluNB subunit and A645 and N616 on the GluN1 subunit, are important for binding of memantine and related compounds as seen in figure 2.<a href="#cite_note-Limapichat-32">[32]</a> The NMDA receptor forms a <a href="/wiki/Heterotetramer" class="mw-redirect" title="Heterotetramer">heterotetramer</a> between two GluN1 and two GluN2 subunits (the subunits were previously denoted as GluN1 and GluN2), two obligatory GluN1 subunits and two regionally localized GluN2 subunits. A related <a href="/wiki/Gene" title="Gene">gene</a> family of GluN3 A and B subunits have an inhibitory effect on receptor activity. Multiple receptor <a href="/wiki/Isoform" class="mw-redirect" title="Isoform">isoforms</a> with distinct brain distributions and functional properties arise by selective splicing of the GluN1 transcripts and differential expression of the GluN2 subunits. Each receptor subunit has modular design and each structural module, also represents a functional unit: <ul><li>The <a href="/wiki/Extracellular" class="mw-redirect" title="Extracellular">extracellular</a> <a href="/wiki/Protein_domain" title="Protein domain">domain</a> contains two globular structures: a modulatory domain and a <a href="/wiki/Ligand" title="Ligand">ligand</a>-binding domain. GluN1 subunits bind the co-agonist glycine and GluN2 subunits bind the neurotransmitter glutamate.<a href="#cite_note-Laube-1">[1]</a><a href="#cite_note-Anson-2">[2]</a></li> <li>The agonist-binding module links to a membrane domain, which consists of three transmembrane segments and a re-entrant loop reminiscent of the selectivity filter of <a href="/wiki/Potassium_channels" class="mw-redirect" title="Potassium channels">potassium channels</a>.</li> <li>The membrane domain contributes residues to the channel pore and is responsible for the receptor's high-unitary <a href="/wiki/Electrical_conductance" class="mw-redirect" title="Electrical conductance">conductance</a>, high-calcium permeability, and voltage-dependent magnesium block.</li> <li>Each subunit has an extensive cytoplasmic domain, which contain residues that can be directly modified by a series of <a href="/wiki/Protein_kinases" class="mw-redirect" title="Protein kinases">protein kinases</a> and <a href="/wiki/Protein_phosphatases" class="mw-redirect" title="Protein phosphatases">protein phosphatases</a>, as well as residues that interact with a large number of structural, adaptor, and scaffolding proteins.</li></ul> The glycine-binding modules of the GluN1 and GluN3 subunits and the glutamate-binding module of the GluN2A subunit have been expressed as soluble proteins, and their three-dimensional structure has been solved at atomic resolution by <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">x-ray crystallography</a>. This has revealed a common fold with amino acid-binding bacterial proteins and with the glutamate-binding module of AMPA-receptors and kainate-receptors. <div class="mw-heading mw-heading2"><h2 id="Mechanism_of_action">Mechanism of action</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=4" title="Edit section: Mechanism of action">edit</a>]</div> NMDA receptors are a crucial part of the development of the central nervous system. The processes of learning, memory, and <a href="/wiki/Neuroplasticity" title="Neuroplasticity">neuroplasticity</a> rely on the mechanism of NMDA receptors. NMDA receptors are glutamate-gated cation channels that allow for an increase of calcium <a href="/wiki/Permeability_(Earth_sciences)" class="mw-redirect" title="Permeability (Earth sciences)">permeability</a>. Channel activation of NMDA receptors is a result of the binding of two co agonists, <a href="/wiki/Glycine" title="Glycine">glycine</a> and <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>. Overactivation of NMDA receptors, causing excessive influx of Ca2+ can lead to excitotoxicity. Excitotoxicity is implied to be involved in some neurodegenerative disorders such as Alzheimer's disease, Parkinson's disease and Huntington's disease.<a href="#cite_note-Chen-16">[16]</a><a href="#cite_note-Kemp-17">[17]</a><a href="#cite_note-Lipton1-18">[18]</a><a href="#cite_note-Koch-19">[19]</a> Blocking of NMDA receptors could therefore, in theory, be useful in treating such diseases.<a href="#cite_note-Chen-16">[16]</a><a href="#cite_note-Kemp-17">[17]</a><a href="#cite_note-Lipton1-18">[18]</a> It is, however, important to preserve physiological NMDA receptor activity while trying to block its excessive, excitotoxic activity. This can possibly be achieved by uncompetitive antagonists, blocking the receptors ion channel when excessively open.<a href="#cite_note-Lipton1-18">[18]</a> Uncompetitive NMDA receptor antagonists, or channel blockers, enter the channel of the NMDA receptor after it has been activated and thereby block the flow of ions.<a href="#cite_note-Johnson-14">[14]</a><a href="#cite_note-Chen-16">[16]</a> <a href="/wiki/MK-801" class="mw-redirect" title="MK-801">MK-801</a>, <a href="/wiki/Ketamine" title="Ketamine">ketamine</a>, <a href="/wiki/Amantadine" title="Amantadine">amantadine</a> and <a href="/wiki/Memantine" title="Memantine">memantine</a> are examples of such antagonists,<a href="#cite_note-Johnson-14">[14]</a> see figure 1. The off-rate of an antagonist from the receptors channel is an important factor as too slow off-rate can interfere with normal function of the receptor and too fast off-rate may give ineffective blockade of an excessively open receptor.<a href="#cite_note-Lipton1-18">[18]</a> <a href="/wiki/Memantine" title="Memantine">Memantine</a> is an example of an uncompetitive channel blocker of the NMDA receptor, with a relatively rapid off-rate and low affinity. At physiological pH its amine group is positively charged and its receptor antagonism is voltage-dependent.<a href="#cite_note-Lipton1-18">[18]</a> It thereby mimics the physiological function of Mg2+ as channel blocker.<a href="#cite_note-Dominguez-15">[15]</a> Memantine only blocks NMDA receptor associated channels during prolonged activation of the receptor, as it occurs under excitotoxic conditions, by replacing magnesium at the binding site. During normal receptor activity the channels only stay open for several milliseconds and under those circumstances memantine is unable to bind within the channels and therefore does not interfere with normal synaptic activity.<a href="#cite_note-Lipton2-21">[21]</a> <div class="mw-heading mw-heading2"><h2 id="Variants">Variants</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=5" title="Edit section: Variants">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="GluN1">GluN1</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=6" title="Edit section: GluN1">edit</a>]</div> There are eight variants of the <a href="/wiki/GRIN1" title="GRIN1">GluN1</a> subunit produced by alternative splicing of <a href="/wiki/GRIN1" title="GRIN1">GRIN1</a>:<a href="#cite_note-Stephenson_2006-35">[35]</a> <ul><li>GluN1-1a, GluN1-1b; GluN1-1a is the most abundantly expressed form.</li> <li>GluN1-2a, GluN1-2b;</li> <li>GluN1-3a, GluN1-3b;</li> <li>GluN1-4a, GluN1-4b;</li></ul> <div class="mw-heading mw-heading3"><h3 id="GluN2">GluN2</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=7" title="Edit section: GluN2">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Model_of_NR2_Subunit_of_NMDA_receptor_(vertebrate_and_invertebrate).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg/220px-Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg/330px-Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg/440px-Model_of_NR2_Subunit_of_NMDA_receptor_%28vertebrate_and_invertebrate%29.jpg 2x" data-file-width="631" data-file-height="472" /></a><figcaption>NR2 subunit in vertebrates (left) and invertebrates (right). Ryan et al., 2008</figcaption></figure> While a single GluN2 subunit is found in <a href="/wiki/Invertebrate" title="Invertebrate">invertebrate</a> <a href="/wiki/Organism" title="Organism">organisms</a>, four distinct isoforms of the GluN2 subunit are expressed in <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> and are referred to with the nomenclature GluN2A through GluN2D (encoded by <a href="/wiki/GRIN2A" title="GRIN2A">GRIN2A</a>, <a href="/wiki/GRIN2B" title="GRIN2B">GRIN2B</a>, <a href="/wiki/GRIN2C" title="GRIN2C">GRIN2C</a>, <a href="/wiki/GRIN2D" title="GRIN2D">GRIN2D</a>). Strong evidence shows that the genes encoding the GluN2 subunits in vertebrates have undergone at least two rounds of <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a>.<a href="#cite_note-pmid20976280-36">[36]</a> They contain the binding-site for <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>. More importantly, each GluN2 subunit has a different intracellular C-terminal domain that can interact with different sets of signaling molecules.<a href="#cite_note-Ryan2009-37">[37]</a> Unlike GluN1 subunits, GluN2 subunits are expressed differentially across various cell types and developmental timepoints and control the electrophysiological properties of the NMDA receptor. In classic circuits, GluN2B is mainly present in immature neurons and in extrasynaptic locations such as <a href="/wiki/Growth_cone" title="Growth cone">growth cones</a>,<a href="#cite_note-Georgiev2008-38">[38]</a> and contains the binding-site for the selective inhibitor <a href="/wiki/Ifenprodil" title="Ifenprodil">ifenprodil</a>.<a href="#cite_note-Bunk2014-39">[39]</a> However, in <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal cell</a> <a href="/wiki/Synapse" title="Synapse">synapses</a> in the newly evolved primate <a href="/wiki/Dorsolateral_prefrontal_cortex" title="Dorsolateral prefrontal cortex">dorsolateral prefrontal cortex</a>, GluN2B are exclusively within the <a href="/wiki/Postsynaptic_density" title="Postsynaptic density">postsynaptic density</a>, and mediate higher cognitive operations such as <a href="/wiki/Working_memory" title="Working memory">working memory</a>.<a href="#cite_note-Wang2013-40">[40]</a> This is consistent with the expansion in GluN2B actions and expression across the cortical hierarchy in <a href="/wiki/Monkey" title="Monkey">monkeys</a> <a href="#cite_note-Yang2018-41">[41]</a> and <a href="/wiki/Human" title="Human">humans</a> <a href="#cite_note-Burt2018-42">[42]</a> and across <a href="/wiki/Primate" title="Primate">primate</a> <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cortex</a> <a href="/wiki/Evolution" title="Evolution">evolution</a>.<a href="#cite_note-Muntane2015-43">[43]</a> <div class="mw-heading mw-heading3"><h3 id="GluN2B_to_GluN2A_switch">GluN2B to GluN2A switch</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=8" title="Edit section: GluN2B to GluN2A switch">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:NR2B-NR2A_switch_in_human_cerebellum,_microarrays,_Bar-Shira_et_al_2015.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/8/8c/NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png/220px-NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png" decoding="async" width="220" height="161" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8c/NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png/330px-NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8c/NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png/440px-NR2B-NR2A_switch_in_human_cerebellum%2C_microarrays%2C_Bar-Shira_et_al_2015.png 2x" data-file-width="742" data-file-height="542" /></a><figcaption>The timecourse of GluN2B-GluN2A switch in human cerebellum. Bar-Shira et al., 2015 <a href="#cite_note-pmid26636753-44">[44]</a></figcaption></figure> While <a href="/wiki/GRIN2B" title="GRIN2B">GluN2B</a> is predominant in the early postnatal brain, the number of GluN2A subunits increases during early development; eventually, <a href="/wiki/GRIN2A" title="GRIN2A">GluN2A</a> subunits become more numerous than GluN2B. This is called the GluN2B-GluN2A developmental switch, and is notable because of the different kinetics each GluN2 subunit contributes to receptor function.<a href="#cite_note-pmid15470155-45">[45]</a> For instance, greater ratios of the GluN2B subunit leads to NMDA receptors which remain open longer compared to those with more GluN2A.<a href="#cite_note-pmid10789248-46">[46]</a> This may in part account for greater memory abilities in the immediate postnatal period compared to late in life, which is the principle behind genetically altered '<a href="/wiki/Doogie_mice" class="mw-redirect" title="Doogie mice">doogie mice</a>'. The detailed time course of this switch in the human cerebellum has been estimated using expression microarray and RNA seq and is shown in the figure on the right. There are three hypothetical models to describe this switch mechanism: <ul><li>Increase in synaptic GluN2A along with decrease in GluN2B</li> <li>Extrasynaptic displacement of GluN2B away from the synapse with increase in GluN2A</li> <li>Increase of GluN2A diluting the number of GluN2B without the decrease of the latter.</li></ul> The GluN2B and GluN2A subunits also have differential roles in mediating <a href="/wiki/Excitotoxicity" title="Excitotoxicity">excitotoxic</a> neuronal death.<a href="#cite_note-pmid17360906-47">[47]</a> The developmental switch in subunit composition is thought to explain the developmental changes in NMDA neurotoxicity.<a href="#cite_note-pmid16540573-48">[48]</a> Homozygous disruption of the gene for GluN2B in mice causes perinatal <a href="/wiki/Lethality" title="Lethality">lethality</a>, whereas disruption of the GluN2A gene produces viable mice, although with impaired hippocampal plasticity.<a href="#cite_note-49">[49]</a> One study suggests that <a href="/wiki/Reelin" title="Reelin">reelin</a> may play a role in the NMDA receptor maturation by increasing the GluN2B subunit mobility.<a href="#cite_note-pmid17881522-50">[50]</a> <div class="mw-heading mw-heading3"><h3 id="GluN2B_to_GluN2C_switch">GluN2B to GluN2C switch</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=9" title="Edit section: GluN2B to GluN2C switch">edit</a>]</div> Granule cell precursors (GCPs) of the cerebellum, after undergoing symmetric cell division<a href="#cite_note-pmid18322077-51">[51]</a> in the external granule-cell layer (EGL), migrate into the internal granule-cell layer (IGL) where they down-regulate GluN2B and activate GluN2C, a process that is independent of neuregulin beta signaling through ErbB2 and ErbB4 receptors.<a href="#cite_note-pmid19244516-52">[52]</a> <div class="mw-heading mw-heading2"><h2 id="Role_in_excitotoxicity">Role in excitotoxicity</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=10" title="Edit section: Role in excitotoxicity">edit</a>]</div> NMDA receptors have been implicated by a number of studies to be strongly involved with <a href="/wiki/Excitotoxicity" title="Excitotoxicity">excitotoxicity</a>.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-pmid2892896-54">[54]</a><a href="#cite_note-55">[55]</a> Because NMDA receptors play an important role in the health and function of <a href="/wiki/Neuron" title="Neuron">neurons</a>, there has been much discussion on how these receptors can affect both cell survival and cell death.<a href="#cite_note-:2-56">[56]</a> Recent evidence supports the hypothesis that overstimulation of <a href="/wiki/Extrasynaptic_NMDA_receptor" title="Extrasynaptic NMDA receptor">extrasynaptic NMDA receptors</a> has more to do with excitotoxicity than stimulation of their <a href="/wiki/Synapse" title="Synapse">synaptic</a> counterparts.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-pmid11953750-24">[24]</a> In addition, while stimulation of <a href="/wiki/Extrasynaptic_NMDA_receptor" title="Extrasynaptic NMDA receptor">extrasynaptic NMDA receptors</a> appear to contribute to cell death, there is evidence to suggest that stimulation of synaptic NMDA receptors contributes to the health and longevity of the cell. There is ample evidence to support the dual nature of NMDA receptors based on location, and the hypothesis explaining the two differing mechanisms is known as the "localization hypothesis".<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-:2-56">[56]</a> <div class="mw-heading mw-heading3"><h3 id="Differing_cascade_pathways">Differing cascade pathways</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=11" title="Edit section: Differing cascade pathways">edit</a>]</div> In order to support the localization hypothesis, it would be necessary to show differing <a href="/wiki/Cell_signaling" title="Cell signaling">cellular signaling pathways</a> are activated by NMDA receptors based on its location within the cell membrane.<a href="#cite_note-:0-53">[53]</a> Experiments have been designed to stimulate either synaptic or non-synaptic NMDA receptors exclusively. These types of experiments have shown that different pathways are being activated or regulated depending on the location of the signal origin.<a href="#cite_note-pmid20720132-57">[57]</a> Many of these pathways use the same <a href="/wiki/Protein_targeting" title="Protein targeting">protein signals</a>, but are regulated oppositely by NMDARs depending on its location. For example, synaptic NMDA excitation caused a decrease in the intracellular concentration of p38 mitogen-activated protein kinase (<a href="/wiki/P38_mitogen-activated_protein_kinases" title="P38 mitogen-activated protein kinases">p38MAPK</a>). Extrasynaptic stimulation NMDARs regulated p38MAPK in the opposite fashion, causing an increase in intracellular concentration.<a href="#cite_note-WangBriz2013-58">[58]</a><a href="#cite_note-pmid19625523-59">[59]</a> Experiments of this type have since been repeated with the results indicating these differences stretch across many pathways linked to cell survival and excitotoxicity.<a href="#cite_note-:0-53">[53]</a> Two specific proteins have been identified as a major pathway responsible for these different cellular responses <a href="/wiki/Extracellular_signal-regulated_kinases" title="Extracellular signal-regulated kinases">ERK1/2</a>, and Jacob.<a href="#cite_note-:0-53">[53]</a> ERK1/2 is responsible for phosphorylation of Jacob when excited by synaptic NMDARs. This information is then <a href="/wiki/Nuclear_transport" title="Nuclear transport">transported to the nucleus</a>. Phosphorylation of Jacob does not take place with extrasynaptic NMDA stimulation. This allows the <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> in the nucleus to respond differently based in the phosphorylation state of Jacob.<a href="#cite_note-KarpovaMikhaylova2013-60">[60]</a> <div class="mw-heading mw-heading3"><h3 id="Neural_plasticity">Neural plasticity</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=12" title="Edit section: Neural plasticity">edit</a>]</div> NMDA receptors (NMDARs) critically influence the induction of synaptic plasticity. NMDARs trigger both long-term potentiation (LTP) and long-term depression (LTD) via fast synaptic transmission.<a href="#cite_note-61">[61]</a> Experimental data suggest that extrasynaptic NMDA receptors inhibit LTP while producing LTD.<a href="#cite_note-pmid21543591-62">[62]</a> Inhibition of LTP can be prevented with the introduction of a <a href="/wiki/NMDA_receptor_antagonist" title="NMDA receptor antagonist">NMDA antagonist</a>.<a href="#cite_note-:0-53">[53]</a> A <a href="/wiki/Transcranial_magnetic_stimulation" title="Transcranial magnetic stimulation">theta burst stimulation</a> that usually induces LTP with synaptic NMDARs, when applied selectively to extrasynaptic NMDARs produces a LTD.<a href="#cite_note-LiuYang2013-63">[63]</a> Experimentation also indicates that extrasynaptic activity is not required for the formation of LTP. In addition, both synaptic and extrasynaptic activity are involved in expressing a full LTD.<a href="#cite_note-PapouinLadépêche2012-64">[64]</a> <div class="mw-heading mw-heading3"><h3 id="Role_of_differing_subunits">Role of differing subunits</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=13" title="Edit section: Role of differing subunits">edit</a>]</div> Another factor that seems to affect NMDAR induced toxicity is the observed variation in <a href="/wiki/Protein_subunit" title="Protein subunit">subunit</a> makeup. NMDA receptors are heterotetramers with two GluN1 subunits and two variable subunits.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-65">[65]</a> Two of these variable subunits, GluN2A and GluN2B, have been shown to preferentially lead to cell survival and cell death cascades respectively. Although both subunits are found in synaptic and extrasynaptic NMDARs there is some evidence to suggest that the GluN2B subunit occurs more frequently in extrasynaptic receptors. This observation could help explain the dualistic role that NMDA receptors play in excitotoxicity.<a href="#cite_note-pmid20096331-66">[66]</a><a href="#cite_note-pmid21310659-67">[67]</a> t-NMDA receptors have been implicated in excitotoxicity-mediated death of neurons in <a href="/wiki/Temporal_lobe_epilepsy" title="Temporal lobe epilepsy">temporal lobe epilepsy</a>.<a href="#cite_note-:3-68">[68]</a> Despite the compelling evidence and the relative simplicity of these two theories working in tandem, there is still disagreement about the significance of these claims. Some problems in proving these theories arise with the difficulty of using pharmacological means to determine the subtypes of specific NMDARs.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-:1-69">[69]</a> In addition, the theory of subunit variation does not explain how this effect might predominate, as it is widely held that the most common tetramer, made from two GluN1 subunits and one of each subunit GluN2A and GluN2B, makes up a high percentage of the NMDARs.<a href="#cite_note-:0-53">[53]</a> The subunit composition of t-NMDA receptors has recently been visualized in brain tissue.<a href="#cite_note-70">[70]</a> <div class="mw-heading mw-heading3"><h3 id="Excitotoxicity_in_a_clinical_setting">Excitotoxicity in a clinical setting</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=14" title="Edit section: Excitotoxicity in a clinical setting">edit</a>]</div> Excitotoxicity has been thought to play a role in the degenerative properties of <a href="/wiki/Neurodegeneration" class="mw-redirect" title="Neurodegeneration">neurodegenerative</a> conditions since the late 1950s.<a href="#cite_note-71">[71]</a> NMDA receptors seem to play an important role in many of these degenerative diseases affecting the brain. Most notably, excitotoxic events involving NMDA receptors have been linked to Alzheimer's disease and Huntington's disease, as well as with other medical conditions such as strokes and epilepsy.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-pmid20152125-72">[72]</a> Treating these conditions with one of the many known NMDA receptor antagonists, however, leads to a variety of unwanted side effects, some of which can be severe. These side effects are, in part, observed because the NMDA receptors do not just signal for cell death but also play an important role in its vitality.<a href="#cite_note-:2-56">[56]</a> Treatment for these conditions might be found in blocking NMDA receptors not found at the synapse.<a href="#cite_note-:0-53">[53]</a><a href="#cite_note-:4-25">[25]</a> One class of excitotoxicity in disease includes gain-of-function mutations in GRIN2B and GRIN1 associated with cortical malformations, such as <a href="/wiki/Polymicrogyria" title="Polymicrogyria">polymicrogyria</a>.<a href="#cite_note-73">[73]</a> D-serine, an antagonist/inverse co-agonist of t-NMDA receptors, which is made in the brain, has been shown to mitigate neuron loss in an animal model of <a href="/wiki/Temporal_lobe_epilepsy" title="Temporal lobe epilepsy">temporal lobe epilepsy</a>.<a href="#cite_note-:3-68">[68]</a> <div class="mw-heading mw-heading2"><h2 id="Ligands">Ligands</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=15" title="Edit section: Ligands">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Agonists">Agonists</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=16" title="Edit section: Agonists">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg/200px-L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg.png" decoding="async" width="200" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg/300px-L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg/400px-L-Glutamins%C3%A4ure_-_L-Glutamic_acid.svg.png 2x" data-file-width="256" data-file-height="121" /></a><figcaption>L-<a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> (glutamate), the major endogenous agonist of the main site of the NMDAR</figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Glycine-2D-skeletal.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Glycine-2D-skeletal.svg/150px-Glycine-2D-skeletal.svg.png" decoding="async" width="150" height="86" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Glycine-2D-skeletal.svg/225px-Glycine-2D-skeletal.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Glycine-2D-skeletal.svg/300px-Glycine-2D-skeletal.svg.png 2x" data-file-width="1100" data-file-height="631" /></a><figcaption><a href="/wiki/Glycine" title="Glycine">Glycine</a>, the major endogenous agonist of the glycine co-agonist site of the NMDAR</figcaption></figure> Activation of NMDA receptors requires binding of <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamate</a> or <a href="/wiki/Aspartic_acid" title="Aspartic acid">aspartate</a> (aspartate does not stimulate the receptors as strongly).<a href="#cite_note-pmid15703381-74">[74]</a> In addition, NMDARs also require the binding of the <a href="/wiki/Co-agonist" class="mw-redirect" title="Co-agonist">co-agonist</a> <a href="/wiki/Glycine" title="Glycine">glycine</a> for the efficient opening of the ion channel, which is a part of this receptor. <a href="/wiki/D-Serine" class="mw-redirect" title="D-Serine">D-Serine</a> has also been found to co-agonize the NMDA receptor with even greater potency than glycine.<a href="#cite_note-pmid17033043-75">[75]</a> It is produced by <a href="/wiki/Serine_racemase" title="Serine racemase">serine racemase</a>, and is enriched in the same areas as NMDA receptors. Removal of D-serine can block NMDA-mediated excitatory neurotransmission in many areas. Recently, it has been shown that D-serine can be released both by neurons and astrocytes to regulate NMDA receptors. Note that D-serine has also been shown to work as an antagonist / inverse co-agonist for t-NMDA receptors.<a href="#cite_note-:6-76">[76]</a><a href="#cite_note-:3-68">[68]</a> NMDA receptor (NMDAR)-mediated currents are directly related to membrane depolarization. NMDA agonists therefore exhibit fast <a href="/wiki/Mg_ion_(physiology)" class="mw-redirect" title="Mg ion (physiology)">Mg2+</a> unbinding kinetics, increasing channel open probability with depolarization. This property is fundamental to the role of the NMDA receptor in <a href="/wiki/Memory" title="Memory">memory</a> and <a href="/wiki/Learning" title="Learning">learning</a>, and it has been suggested that this channel is a biochemical substrate of <a href="/wiki/Hebbian_learning" class="mw-redirect" title="Hebbian learning">Hebbian learning</a>, where it can act as a coincidence detector for membrane depolarization and synaptic transmission. <div class="mw-heading mw-heading4"><h4 id="Examples">Examples</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=17" title="Edit section: Examples">edit</a>]</div> Some known NMDA receptor agonists include: <ul><li><a href="/wiki/Amino_acid" title="Amino acid">Amino acids</a> and amino acid derivatives <ul><li><a href="/wiki/Aspartic_acid" title="Aspartic acid">Aspartic acid</a> (aspartate) (<a href="/wiki/Aspartic_acid" title="Aspartic acid">D-aspartic acid</a>, <a href="/wiki/Aspartic_acid" title="Aspartic acid">L-aspartic acid</a>) – endogenous glutamate site agonist. The word N-methyl-D-aspartate (NMDA) is partially derived from D-aspartate.</li> <li><a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> (glutamate) – endogenous glutamate site agonist <ul><li><a href="/wiki/Tetrazolylglycine" title="Tetrazolylglycine">Tetrazolylglycine</a> – synthetic glutamate site agonist</li> <li><a href="/wiki/Homocysteic_acid" title="Homocysteic acid">Homocysteic acid</a> – endogenous glutamate site agonist</li> <li><a href="/wiki/Ibotenic_acid" title="Ibotenic acid">Ibotenic acid</a> – naturally occurring glutamate site agonist found in <a href="/wiki/Amanita_muscaria" title="Amanita muscaria">Amanita muscaria</a></li> <li><a href="/wiki/Quinolinic_acid" title="Quinolinic acid">Quinolinic acid</a> (quinolinate) – endogenous glutamate site agonist</li></ul></li> <li><a href="/wiki/Glycine" title="Glycine">Glycine</a> – endogenous glycine site agonist <ul><li><a href="/wiki/Alanine" title="Alanine">Alanine</a> (<a href="/wiki/Alanine" title="Alanine">D-alanine</a>, <a href="/wiki/Alanine" title="Alanine">L-alanine</a>) – endogenous glycine site agonist</li> <li><a href="/wiki/Milacemide" title="Milacemide">Milacemide</a> – synthetic glycine site agonist; prodrug of <a href="/wiki/Glycine" title="Glycine">glycine</a></li> <li><a href="/wiki/Sarcosine" title="Sarcosine">Sarcosine</a> (monomethylglycine) – endogenous glycine site agonist</li> <li><a href="/wiki/Serine" title="Serine">Serine</a> (<a href="/wiki/Serine" title="Serine">D-serine</a>, <a href="/wiki/Serine" title="Serine">L-serine</a>) – endogenous glycine site agonist</li></ul></li></ul></li> <li><a href="/wiki/Positive_allosteric_modulator" class="mw-redirect" title="Positive allosteric modulator">Positive allosteric modulators</a> <ul><li><a href="/wiki/Cerebrosterol" class="mw-redirect" title="Cerebrosterol">Cerebrosterol</a> – endogenous weak positive allosteric modulator</li> <li><a href="/wiki/Cholesterol" title="Cholesterol">Cholesterol</a> – endogenous weak positive allosteric modulator</li> <li><a href="/wiki/Dehydroepiandrosterone" title="Dehydroepiandrosterone">Dehydroepiandrosterone</a> (DHEA) – endogenous weak positive allosteric modulator</li> <li><a href="/wiki/Dehydroepiandrosterone_sulfate" title="Dehydroepiandrosterone sulfate">Dehydroepiandrosterone sulfate</a> (DHEA-S) – endogenous weak positive allosteric modulator</li> <li><a href="/wiki/Nebostinel" class="mw-redirect" title="Nebostinel">Nebostinel</a> (neboglamine) – synthetic positive allosteric modulator of the glycine site</li> <li><a href="/wiki/Pregnenolone_sulfate" title="Pregnenolone sulfate">Pregnenolone sulfate</a> – endogenous weak positive allosteric modulator</li></ul></li> <li>Polyamines <ul><li><a href="/wiki/Spermidine" title="Spermidine">Spermidine</a> – endogenous polyamine site agonist</li> <li><a href="/wiki/Spermine" title="Spermine">Spermine</a> – endogenous polyamine site agonist</li></ul></li></ul> <div class="mw-heading mw-heading4"><h4 id="Neramexane">Neramexane</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=18" title="Edit section: Neramexane">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Neramexane.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Neramexane.svg/220px-Neramexane.svg.png" decoding="async" width="220" height="202" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Neramexane.svg/330px-Neramexane.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/95/Neramexane.svg/440px-Neramexane.svg.png 2x" data-file-width="512" data-file-height="471" /></a><figcaption>Figure 6: Chemical structure of neramexane, second generation memantine derivative</figcaption></figure> An example of memantine derivative is <a href="/wiki/Neramexane" title="Neramexane">neramexane</a> which was discovered by studying number of aminoalkyl <a href="/wiki/Cyclohexanes" class="mw-redirect" title="Cyclohexanes">cyclohexanes</a>, with memantine as the template, as NMDA receptor antagonists. Neramexane binds to the same site as memantine within the NMDA receptor associated channel and with comparable affinity. It does also show very similar bioavailability and blocking kinetics <a href="/wiki/In_vivo" title="In vivo">in vivo</a> as memantine. Neramexane went to <a href="/wiki/Clinical_trials" class="mw-redirect" title="Clinical trials">clinical trials</a> for four indications, including Alzheimer's disease.<a href="#cite_note-Wanka-28">[28]</a> <div class="mw-heading mw-heading3"><h3 id="Partial_agonists">Partial agonists</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=19" title="Edit section: Partial agonists">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:NMDA.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/NMDA.svg/200px-NMDA.svg.png" decoding="async" width="200" height="123" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/NMDA.svg/300px-NMDA.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b9/NMDA.svg/400px-NMDA.svg.png 2x" data-file-width="446" data-file-height="274" /></a><figcaption><a href="/wiki/N-Methyl-D-aspartic_acid" title="N-Methyl-D-aspartic acid">N-Methyl-D-aspartic acid</a> (NMDA), a synthetic partial agonist of the main site of the NMDAR</figcaption></figure> <a href="/wiki/N-Methyl-D-aspartic_acid" title="N-Methyl-D-aspartic acid">N-Methyl-D-aspartic acid</a> (NMDA), which the NMDA receptor was named after, is a partial agonist of the active or glutamate recognition site. 3,5-Dibromo-L-phenylalanine, a naturally occurring halogenated derivative of <a href="/wiki/Phenylalanine" title="Phenylalanine">L-phenylalanine</a>, is a weak partial NMDA receptor agonist acting on the glycine site.<a href="#cite_note-77">[77]</a> 3,5-Dibromo-L-phenylalanine has been proposed a novel therapeutic drug candidate for treatment of neuropsychiatric disorders and diseases such as <a href="/wiki/Schizophrenia" title="Schizophrenia">schizophrenia</a>,<a href="#cite_note-78">[78]</a> and neurological disorders such as <a href="/wiki/Ischemic_stroke" class="mw-redirect" title="Ischemic stroke">ischemic stroke</a> and <a href="/wiki/Epileptic_seizure" class="mw-redirect" title="Epileptic seizure">epileptic seizures</a>.<a href="#cite_note-79">[79]</a> Other partial agonists of the NMDA receptor acting on novel sites such as <a href="/wiki/Rapastinel" title="Rapastinel">rapastinel</a> (GLYX-13) and <a href="/wiki/Apimostinel" title="Apimostinel">apimostinel</a> (NRX-1074) are now viewed for the development of new drugs with antidepressant and analgesic effects without obvious psychotomimetic activities.<a href="#cite_note-80">[80]</a> <div class="mw-heading mw-heading4"><h4 id="Examples_2">Examples</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=20" title="Edit section: Examples">edit</a>]</div> <ul><li><a href="/wiki/Aminocyclopropanecarboxylic_acid" class="mw-redirect" title="Aminocyclopropanecarboxylic acid">Aminocyclopropanecarboxylic acid</a> (ACC) – synthetic glycine site partial agonist</li> <li><a href="/wiki/Cycloserine" title="Cycloserine">Cycloserine</a> (<a href="/wiki/D-cycloserine" class="mw-redirect" title="D-cycloserine">D-cycloserine</a>) – naturally occurring glycine site partial agonist found in <a href="/wiki/Streptomyces" title="Streptomyces">Streptomyces orchidaceus</a></li> <li><a href="/wiki/HA-966" title="HA-966">HA-966</a> – synthetic glycine site weak partial agonist</li> <li><a href="/wiki/Homoquinolinic_acid" title="Homoquinolinic acid">Homoquinolinic acid</a> – synthetic glutamate site partial agonist</li> <li><a href="/wiki/N-Methyl-D-aspartic_acid" title="N-Methyl-D-aspartic acid">N-Methyl-D-aspartic acid</a> (NMDA) – synthetic glutamate site partial agonist</li></ul> Positive allosteric modulators include: <ul><li><a href="/wiki/AGN-241751" class="mw-redirect" title="AGN-241751">Zelquistinel</a> (GATE-251) – synthetic novel site partial agonist</li> <li><a href="/wiki/Apimostinel" title="Apimostinel">Apimostinel</a> (GATE-202) – synthetic novel site partial agonist</li> <li><a href="/wiki/Rapastinel" title="Rapastinel">Rapastinel</a> (GLYX-13) – synthetic novel site partial agonist<a href="#cite_note-81">[81]</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Antagonists">Antagonists</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=21" title="Edit section: Antagonists">edit</a>]</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/NMDA_receptor_antagonist" title="NMDA receptor antagonist">NMDA receptor antagonist</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Ketamine.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Ketamine.svg/150px-Ketamine.svg.png" decoding="async" width="150" height="176" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Ketamine.svg/225px-Ketamine.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/56/Ketamine.svg/300px-Ketamine.svg.png 2x" data-file-width="580" data-file-height="680" /></a><figcaption><a href="/wiki/Ketamine" title="Ketamine">Ketamine</a>, a synthetic general anesthetic and one of the best-known NMDAR antagonists</figcaption></figure> Antagonists of the NMDA receptor are used as <a href="/wiki/Anesthetic" title="Anesthetic">anesthetics</a> for animals and sometimes humans, and are often used as <a href="/wiki/Recreational_drug" class="mw-redirect" title="Recreational drug">recreational drugs</a> due to their <a href="/wiki/Hallucinogenic" class="mw-redirect" title="Hallucinogenic">hallucinogenic</a> properties, in addition to their unique effects at elevated dosages such as <a href="/wiki/Dissociation_(psychology)" title="Dissociation (psychology)">dissociation</a>. When certain NMDA receptor antagonists are given to rodents in large doses, they can cause a form of <a href="/wiki/Brain_damage" class="mw-redirect" title="Brain damage">brain damage</a> called <a href="/wiki/Olney%27s_lesions" title="Olney's lesions">Olney's lesions</a>. NMDA receptor antagonists that have been shown to induce Olney's lesions include <a href="/wiki/Ketamine" title="Ketamine">ketamine</a>, <a href="/wiki/Phencyclidine" title="Phencyclidine">phencyclidine</a>, and <a href="/wiki/Dextrorphan" title="Dextrorphan">dextrorphan</a> (a metabolite of <a href="/wiki/Dextromethorphan" title="Dextromethorphan">dextromethorphan</a>), as well as some NMDA receptor antagonists used only in research environments. So far, the published research on Olney's lesions is inconclusive in its occurrence upon human or monkey brain tissues with respect to an increase in the presence of NMDA receptor antagonists.<a href="#cite_note-urlErowid_DXM_Vaults-82">[82]</a> Most NMDAR antagonists are <a href="/wiki/Uncompetitive_antagonist" class="mw-redirect" title="Uncompetitive antagonist">uncompetitive</a> or <a href="/wiki/Noncompetitive_antagonist" class="mw-redirect" title="Noncompetitive antagonist">noncompetitive blockers</a> of the channel pore or are antagonists of the glycine co-regulatory site rather than antagonists of the active/glutamate site. <div class="mw-heading mw-heading4"><h4 id="Examples_3">Examples</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=22" title="Edit section: Examples">edit</a>]</div> Common agents in which NMDA receptor antagonism is the primary or a major mechanism of action: <ul><li><a href="/wiki/4-Chlorokynurenine" title="4-Chlorokynurenine">4-Chlorokynurenine</a> (AV-101) – glycine site antagonist; prodrug of <a href="/wiki/7-chlorokynurenic_acid" class="mw-redirect" title="7-chlorokynurenic acid">7-chlorokynurenic acid</a><a href="#cite_note-Flight2013-83">[83]</a><a href="#cite_note-VécseiSzalárdy2012-84">[84]</a></li> <li><a href="/wiki/7-Chlorokynurenic_acid" title="7-Chlorokynurenic acid">7-Chlorokynurenic acid</a> – glycine site antagonist</li> <li><a href="/wiki/Agmatine" title="Agmatine">Agmatine</a> – endogenous polyamine site antagonist<a href="#cite_note-pmid10785653-85">[85]</a><a href="#cite_note-pmid12363406-86">[86]</a></li> <li><a href="/w/index.php?title=Argiotoxin-636&action=edit&redlink=1" class="new" title="Argiotoxin-636 (page does not exist)">Argiotoxin-636</a> – naturally occurring dizocilpine or related site antagonist found in <a href="/wiki/Argiope_(spider)" title="Argiope (spider)">Argiope</a> venom</li> <li><a href="/wiki/AP5" title="AP5">AP5</a> – glutamate site antagonist</li> <li><a href="/wiki/AP-7_(drug)" title="AP-7 (drug)">AP7</a> – glutamate site antagonist</li> <li><a href="/wiki/CGP-37849" title="CGP-37849">CGP-37849</a> – glutamate site antagonist</li> <li><a href="/wiki/Serine" title="Serine">D-serine</a> - t-NMDA receptor antagonist / inverse co-agonist<a href="#cite_note-:6-76">[76]</a><a href="#cite_note-:3-68">[68]</a></li> <li><a href="/wiki/Delucemine" title="Delucemine">Delucemine</a> (NPS-1506) – dizocilpine or related site antagonist; derived from <a href="/w/index.php?title=Argiotoxin-636&action=edit&redlink=1" class="new" title="Argiotoxin-636 (page does not exist)">argiotoxin-636</a><a href="#cite_note-pmid11026487-87">[87]</a><a href="#cite_note-pmid26257776-88">[88]</a></li> <li><a href="/wiki/Dextromethorphan" title="Dextromethorphan">Dextromethorphan</a> (DXM) – dizocilpine site antagonist; prodrug of <a href="/wiki/Dextrorphan" title="Dextrorphan">dextrorphan</a></li> <li><a href="/wiki/Dextrorphan" title="Dextrorphan">Dextrorphan</a> (DXO) – dizocilpine site antagonist</li> <li><a href="/wiki/Dexanabinol" title="Dexanabinol">Dexanabinol</a> – dizocilpine-related site antagonist<a href="#cite_note-ShohamiMechoulam2000-89">[89]</a><a href="#cite_note-pmid2556719-90">[90]</a><a href="#cite_note-pmid8242387-91">[91]</a></li> <li><a href="/wiki/Diethyl_ether" title="Diethyl ether">Diethyl ether</a> – unknown site antagonist</li> <li><a href="/wiki/Diphenidine" title="Diphenidine">Diphenidine</a> – dizocilpine site antagonist</li> <li><a href="/wiki/Dizocilpine" title="Dizocilpine">Dizocilpine</a> (MK-801) – dizocilpine site antagonist</li> <li><a href="/wiki/Eliprodil" title="Eliprodil">Eliprodil</a> – ifenprodil site antagonist</li> <li><a href="/wiki/Esketamine" title="Esketamine">Esketamine</a> – dizocilpine site antagonist</li> <li><a href="/wiki/Hodgkinsine" title="Hodgkinsine">Hodgkinsine</a> – undefined site antagonist</li> <li><a href="/wiki/Ifenprodil" title="Ifenprodil">Ifenprodil</a> – ifenprodil site antagonist<a href="#cite_note-pmid21677647-92">[92]</a></li> <li><a href="/wiki/Kaitocephalin" title="Kaitocephalin">Kaitocephalin</a> – naturally occurring glutamate site antagonist found in <a href="/wiki/Eupenicillium_shearii" title="Eupenicillium shearii">Eupenicillium shearii</a></li> <li><a href="/wiki/Ketamine" title="Ketamine">Ketamine</a> – dizocilpine site antagonist</li> <li><a href="/wiki/Kynurenic_acid" title="Kynurenic acid">Kynurenic acid</a> – endogenous glycine site antagonist</li> <li><a href="/wiki/Lanicemine" title="Lanicemine">Lanicemine</a> – low-trapping dizocilpine site antagonist</li> <li><a href="/wiki/LY-235959" title="LY-235959">LY-235959</a> – glutamate site antagonist</li> <li><a href="/wiki/Memantine" title="Memantine">Memantine</a> – low-trapping dizocilpine site antagonist</li> <li><a href="/wiki/Methoxetamine" title="Methoxetamine">Methoxetamine</a> – dizocilpine site antagonist</li> <li><a href="/wiki/Midafotel" title="Midafotel">Midafotel</a> – glutamate site antagonist</li> <li><a href="/wiki/Nitrous_oxide" title="Nitrous oxide">Nitrous oxide</a> (N2O) – undefined site antagonist</li> <li><a href="/wiki/PEAQX" title="PEAQX">PEAQX</a> – glutamate site antagonist</li> <li><a href="/wiki/Perzinfotel" title="Perzinfotel">Perzinfotel</a> – glutamate site antagonist</li> <li><a href="/wiki/Phencyclidine" title="Phencyclidine">Phencyclidine</a> (PCP) – dizocilpine site antagonist</li> <li><a href="/wiki/Phenylalanine" title="Phenylalanine">Phenylalanine</a> - a naturally occurring amino acid, glycine site antagonist<a href="#cite_note-pmid11986979-93">[93]</a><a href="#cite_note-94">[94]</a></li> <li><a href="/wiki/Psychotridine" title="Psychotridine">Psychotridine</a> – undefined site antagonist</li> <li><a href="/wiki/Selfotel" title="Selfotel">Selfotel</a> – glutamate site antagonist</li> <li><a href="/wiki/Tiletamine" title="Tiletamine">Tiletamine</a> – dizocilpine site antagonist</li> <li><a href="/wiki/Traxoprodil" title="Traxoprodil">Traxoprodil</a> – ifenprodil site antagonist</li> <li><a href="/wiki/Xenon" title="Xenon">Xenon</a> – unknown site antagonist</li></ul> Some common agents in which weak NMDA receptor antagonism is a secondary or additional action include: <ul><li><a href="/wiki/Amantadine" title="Amantadine">Amantadine</a> – an <a href="/wiki/Antiviral" class="mw-redirect" title="Antiviral">antiviral</a> and <a href="/wiki/Management_of_Parkinson%27s_disease#Medication" title="Management of Parkinson's disease">antiparkinsonian</a> drug; low-trapping dizocilpine site antagonist<a href="#cite_note-95">[95]</a></li> <li><a href="/wiki/Atomoxetine" title="Atomoxetine">Atomoxetine</a> – a <a href="/wiki/Norepinephrine_reuptake_inhibitor" title="Norepinephrine reuptake inhibitor">norepinephrine reuptake inhibitor</a> used to treat <a href="/wiki/Attention-deficit_hyperactivity_disorder" class="mw-redirect" title="Attention-deficit hyperactivity disorder"><abbr title="attention-deficit hyperactivity disorder">ADHD</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip attention-deficit hyperactivity disorder<a href="#cite_note-Atomoxetine_acts_as_an_NMDA_receptor_blocker_in_clinically_relevant_concentrations-96">[96]</a></li> <li><a href="/wiki/Dextropropoxyphene" title="Dextropropoxyphene">Dextropropoxyphene</a> – an <a href="/wiki/Opioid_analgesic" class="mw-redirect" title="Opioid analgesic">opioid analgesic</a></li> <li><a href="/wiki/Ethanol" title="Ethanol">Ethanol</a> (<a href="/wiki/Alcoholic_drink" class="mw-redirect" title="Alcoholic drink">alcohol</a>) – a <a href="/wiki/Euphoriant" class="mw-redirect" title="Euphoriant">euphoriant</a>, <a href="/wiki/Sedative" title="Sedative">sedative</a>, and <a href="/wiki/Anxiolytic" title="Anxiolytic">anxiolytic</a> used recreationally; unknown site antagonist</li> <li><a href="/wiki/Guaifenesin" title="Guaifenesin">Guaifenesin</a> – an <a href="/wiki/Expectorant" class="mw-redirect" title="Expectorant">expectorant</a></li> <li><a href="/wiki/Huperzine_A" title="Huperzine A">Huperzine A</a> – a naturally occurring <a href="/wiki/Acetylcholinesterase_inhibitor" title="Acetylcholinesterase inhibitor">acetylcholinesterase inhibitor</a> and potential <a href="/wiki/Antidementia" class="mw-redirect" title="Antidementia">antidementia</a> agent</li> <li><a href="/wiki/Ibogaine" title="Ibogaine">Ibogaine</a> – a naturally occurring <a href="/wiki/Hallucinogen" title="Hallucinogen">hallucinogen</a> and <a href="/wiki/Antiaddictive" class="mw-redirect" title="Antiaddictive">antiaddictive</a> agent</li> <li><a href="/wiki/Ketobemidone" title="Ketobemidone">Ketobemidone</a> – an opioid analgesic</li> <li><a href="/wiki/Methadone" title="Methadone">Methadone</a> – an opioid analgesic</li> <li><a href="/wiki/Minocycline" title="Minocycline">Minocycline</a> – an <a href="/wiki/Antibiotic" title="Antibiotic">antibiotic</a><a href="#cite_note-pmid28616020-97">[97]</a></li> <li><a href="/wiki/Tramadol" title="Tramadol">Tramadol</a> – an atypical opioid analgesic and <a href="/wiki/Serotonin_releasing_agent" title="Serotonin releasing agent">serotonin releasing agent</a></li></ul> <div class="mw-heading mw-heading4"><h4 id="Nitromemantine">Nitromemantine</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=23" title="Edit section: Nitromemantine">edit</a>]</div> The NMDA receptor is regulated via <a href="/wiki/Nitrosylation" class="mw-redirect" title="Nitrosylation">nitrosylation</a> and aminoadamantane can be used as a target-directed shuttle to bring nitrogen oxide (NO) close to the site within the NMDA receptor where it can nitrosylate and regulate the ion channel conductivity.<a href="#cite_note-Wanka-28">[28]</a> A NO donor that can be used to decrease NMDA receptor activity is the alkyl nitrate nitroglycerin. Unlike many other NO donors, alkyl nitrates do not have potential NO associated <a href="/wiki/Neurotoxic" class="mw-redirect" title="Neurotoxic">neurotoxic</a> effects. Alkyl nitrates donate NO in the form of a nitro group as seen in figure 7, -NO2-, which is a safe donor that avoids neurotoxicity. The nitro group must be targeted to the NMDA receptor, otherwise other effects of NO such as dilatation of blood vessels and consequent <a href="/wiki/Hypotension" title="Hypotension">hypotension</a> could result.<a href="#cite_note-Lipton3-98">[98]</a> <a href="/wiki/Nitromemantine" title="Nitromemantine">Nitromemantine</a> is a second-generation derivative of memantine, it reduces excitotoxicity mediated by overactivation of the glutamatergic system by blocking NMDA receptor without sacrificing safety. Provisional studies in animal models show that nitromemantines are more effective than memantine as neuroprotectants, both <a href="/wiki/In_vitro" title="In vitro">in vitro</a> and in vivo. Memantine and newer derivatives could become very important weapons in the fight against neuronal damage.<a href="#cite_note-Lipton1-18">[18]</a> <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Nitromemantine.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Nitromemantine.jpg/450px-Nitromemantine.jpg" decoding="async" width="450" height="119" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/23/Nitromemantine.jpg 1.5x" data-file-width="457" data-file-height="121" /></a><figcaption>Figure 7: Nitroglycerin donate ONO2 group that leads to second generation memantine analog, nitromemantine</figcaption></figure> <a href="/wiki/Negative_allosteric_modulator" class="mw-redirect" title="Negative allosteric modulator">Negative allosteric modulators</a> include: <ul><li><a href="/wiki/25-Hydroxycholesterol" title="25-Hydroxycholesterol">25-Hydroxycholesterol</a> – endogenous weak negative allosteric modulator</li> <li><a href="/wiki/Conantokin" title="Conantokin">Conantokins</a> – naturally occurring negative allosteric modulators of the polyamine site found in <a href="/wiki/Conus_geographus" title="Conus geographus">Conus geographus</a><a href="#cite_note-pmid1328523-99">[99]</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Modulators">Modulators</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=24" title="Edit section: Modulators">edit</a>]</div> <div class="mw-heading mw-heading4"><h4 id="Examples_4">Examples</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=25" title="Edit section: Examples">edit</a>]</div> The NMDA receptor is modulated by a number of <a href="/wiki/Endogenous" class="mw-redirect" title="Endogenous">endogenous</a> and <a href="/wiki/Exogenous" class="mw-redirect" title="Exogenous">exogenous</a> compounds:<a href="#cite_note-pmid15670959-100">[100]</a> <ul><li><a href="/wiki/Aminoglycoside" title="Aminoglycoside">Aminoglycosides</a> have been shown to have a similar effect to polyamines, and this may explain their neurotoxic effect.</li> <li><a href="/wiki/CDK5" class="mw-redirect" title="CDK5">CDK5</a> regulates the amount of <a href="/wiki/NR2B" class="mw-redirect" title="NR2B">NR2B</a>-containing NMDA receptors on the synaptic membrane, thus affecting <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a>.<a href="#cite_note-pmid17529984-101">[101]</a><a href="#cite_note-pmid18184784-102">[102]</a></li> <li><a href="/wiki/Polyamine" title="Polyamine">Polyamines</a> do not directly activate NMDA receptors, but instead act to potentiate or inhibit glutamate-mediated responses.</li> <li><a href="/wiki/Reelin" title="Reelin">Reelin</a> modulates NMDA function through <a href="/wiki/Src_Family_Kinases" class="mw-redirect" title="Src Family Kinases">Src family kinases</a> and <a href="/wiki/DAB1" title="DAB1">DAB1</a>.<a href="#cite_note-pmid16148228-103">[103]</a> significantly enhancing <a href="/wiki/Long-term_potentiation" title="Long-term potentiation">LTP</a> in the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>.</li> <li><a href="/wiki/Src_(gene)" class="mw-redirect" title="Src (gene)">Src</a> kinase enhances NMDA receptor currents.<a href="#cite_note-pmid9005855-104">[104]</a></li> <li><a href="/wiki/Sodium" title="Sodium">Na+</a>, <a href="/wiki/K_ion_(physiology)" class="mw-redirect" title="K ion (physiology)">K+</a> and <a href="/wiki/Ca_ion_(physiology)" class="mw-redirect" title="Ca ion (physiology)">Ca2+</a> not only pass through the NMDA receptor channel but also modulate the activity of NMDA receptors.<a href="#cite_note-105">[105]</a></li> <li><a href="/wiki/Zinc#Biological_role" title="Zinc">Zn2+</a> and <a href="/wiki/Copper" title="Copper">Cu2+</a> generally block NMDA current activity in a noncompetitive and a voltage-independent manner. However zinc may potentiate or inhibit the current depending on the neural activity.<a href="#cite_note-106">[106]</a></li> <li><a href="/wiki/Lead" title="Lead">Pb</a>2+<a href="#cite_note-107">[107]</a> is a potent NMDAR antagonist. Presynaptic deficits resulting from Pb2+ exposure during synaptogenesis are mediated by disruption of NMDAR-dependent BDNF signaling.</li> <li>Proteins of the <a href="/wiki/Major_histocompatibility_complex" title="Major histocompatibility complex">major histocompatibility complex</a> class I are endogenous negative regulators of NMDAR-mediated currents in the adult hippocampus,<a href="#cite_note-pmid21135233-108">[108]</a> and are required for appropriate NMDAR-induced changes in <a href="/wiki/AMPAR" class="mw-redirect" title="AMPAR">AMPAR</a> trafficking <a href="#cite_note-pmid21135233-108">[108]</a> and NMDAR-dependent <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a> and <a href="/wiki/Learning" title="Learning">learning</a> and <a href="/wiki/Memory" title="Memory">memory</a>.<a href="#cite_note-pmid11118151-109">[109]</a><a href="#cite_note-110">[110]</a></li> <li>The activity of NMDA receptors is also strikingly sensitive to the changes in <a href="/wiki/PH" title="PH">pH</a>, and partially inhibited by the ambient concentration of H+ under physiological conditions.<a href="#cite_note-111">[111]</a> The level of inhibition by H+ is greatly reduced in receptors containing the NR1a subtype, which contains the positively charged insert Exon 5. The effect of this insert may be mimicked by positively charged polyamines and aminoglycosides, explaining their mode of action.</li> <li>NMDA receptor function is also strongly regulated by chemical reduction and oxidation, via the so-called "redox modulatory site."<a href="#cite_note-pmid2696504-112">[112]</a> Through this site, reductants dramatically enhance NMDA channel activity, whereas oxidants either reverse the effects of reductants or depress native responses. It is generally believed that NMDA receptors are modulated by endogenous redox agents such as <a href="/wiki/Glutathione" title="Glutathione">glutathione</a>, <a href="/wiki/Lipoic_acid" title="Lipoic acid">lipoic acid</a>, and the essential nutrient <a href="/wiki/Pyrroloquinoline_quinone" title="Pyrroloquinoline quinone">pyrroloquinoline quinone</a>[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>].</li></ul> <div class="mw-heading mw-heading3"><h3 id="Development_of_NMDA_receptor_antagonists">Development of NMDA receptor antagonists</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=26" title="Edit section: Development of NMDA receptor antagonists">edit</a>]</div> The main problem with the development of NMDA antagonists for neuroprotection is that physiological NMDA receptor activity is essential for normal neuronal function. Complete blockade of all NMDA receptor activity results in side effects such as <a href="/wiki/Hallucinations" class="mw-redirect" title="Hallucinations">hallucinations</a>, agitation and <a href="/wiki/Anesthesia" title="Anesthesia">anesthesia</a>. To be clinically relevant, an NMDA receptor antagonist must limit its action to blockade of excessive activation, without limiting normal function of the receptor.<a href="#cite_note-Lipton2-21">[21]</a> <div class="mw-heading mw-heading4"><h4 id="Competitive_NMDA_receptor_antagonists">Competitive NMDA receptor antagonists</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=27" title="Edit section: Competitive NMDA receptor antagonists">edit</a>]</div> <a href="/wiki/Competitive" class="mw-redirect" title="Competitive">Competitive</a> NMDA receptor antagonists, which were developed first, are not a good option because they compete and bind to the same site (NR2 subunit) on the receptor as the agonist, glutamate, and therefore block normal function also.<a href="#cite_note-Lipton2-21">[21]</a><a href="#cite_note-Monaghan-113">[113]</a> They will block healthy areas of the brain prior to having an impact on pathological areas, because healthy areas contain lower levels of <a href="/wiki/Agonist" title="Agonist">agonist</a> than pathological areas. These antagonists can be displaced from the receptor by high concentration of glutamate which can exist under excitotoxic circumstances.<a href="#cite_note-Chen-16">[16]</a> <div class="mw-heading mw-heading4"><h4 id="Noncompetitive_NMDA_receptor_antagonists">Noncompetitive NMDA receptor antagonists</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=28" title="Edit section: Noncompetitive NMDA receptor antagonists">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:NMDA_receptor_antagonist.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/NMDA_receptor_antagonist.jpg/200px-NMDA_receptor_antagonist.jpg" decoding="async" width="200" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/NMDA_receptor_antagonist.jpg/300px-NMDA_receptor_antagonist.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/8/8d/NMDA_receptor_antagonist.jpg 2x" data-file-width="381" data-file-height="338" /></a><figcaption>Figure 4: The chemical structures of MK-801, phencyclidine and ketamine, high affinity uncompetitive NMDA receptor antagonists</figcaption></figure> Uncompetitive NMDA receptor antagonists block within the ion channel at the Mg2+ site (pore region) and prevent excessive influx of Ca2+. Noncompetitive antagonism refers to a type of block that an increased concentration of glutamate cannot overcome, and is dependent upon prior activation of the receptor by the agonist, i.e. it only enters the channel when it is opened by agonist.<a href="#cite_note-Lipton2-21">[21]</a><a href="#cite_note-Sonkusare-114">[114]</a> <div class="mw-heading mw-heading4"><h4 id="Memantine_and_related_compounds">Memantine and related compounds</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=29" title="Edit section: Memantine and related compounds">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Memantine_and_amantadine.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/3/3e/Memantine_and_amantadine.jpg" decoding="async" width="273" height="160" class="mw-file-element" data-file-width="273" data-file-height="160" /></a><figcaption>Figure 5: Chemical structures of memantine (right) and amantadine (left)</figcaption></figure> Because of these adverse side effects of high affinity blockers, the search for clinically successful NMDA receptor antagonists for neurodegenerative diseases continued and focused on developing low affinity blockers. However the affinity could not be too low and dwell time not too short (as seen with Mg2+) where membrane depolarization relieves the block. The discovery was thereby development of uncompetitive antagonist with longer dwell time than Mg2+ in the channel but shorter than MK-801. That way the drug obtained would only block excessively open NMDA receptor associated channels but not normal neurotransmission.<a href="#cite_note-Lipton2-21">[21]</a><a href="#cite_note-Sonkusare-114">[114]</a> Memantine is that drug. It is a derivative of amantadine which was first an anti-influenza agent but was later discovered by coincidence to have efficacy in Parkinson's disease. Chemical structures of memantine and amantadine can be seen in figure 5. The compound was first thought to be <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> or <a href="/wiki/Anticholinergic" title="Anticholinergic">anticholinergic</a> but was later found to be an NMDA receptor antagonist.<a href="#cite_note-Dominguez-15">[15]</a><a href="#cite_note-Lipton2-21">[21]</a> Memantine is the first drug approved for treatment of severe and more advanced <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a>, which for example anticholinergic drugs do not do much good for.<a href="#cite_note-Sonkusare-114">[114]</a> It helps recovery of synaptic function and in that way improves impaired memory and learning.<a href="#cite_note-Koch-19">[19]</a> In 2015 memantine is also in trials for therapeutic importance in additional neurological disorders.<a href="#cite_note-Lipton3-98">[98]</a> Many second-generation memantine derivatives have been in development that may show even better neuroprotective effects, where the main thought is to use other safe but effective modulatory sites on the NMDA receptor in addition to its associated ion channel.<a href="#cite_note-Lipton3-98">[98]</a> <div class="mw-heading mw-heading3"><h3 id="Structure_activity_relationship_(SAR)">Structure activity relationship (SAR)</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=30" title="Edit section: Structure activity relationship (SAR)">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:SAR_of_amantadine_and_related_compunds.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/1/12/SAR_of_amantadine_and_related_compunds.jpg" decoding="async" width="291" height="242" class="mw-file-element" data-file-width="291" data-file-height="242" /></a><figcaption>Figure 8: Structure activity relationship (SAR) of amantadine and related compounds</figcaption></figure> Memantine (1-amino-3,5-dimethyladamantane) is an aminoalkyl cyclohexane derivative and an atypical drug compound with non-planar, three dimensional tricyclic structure. Figure 8 shows SAR for aminoalkyl cyclohexane derivative. Memantine has several important features in its structure for its effectiveness: <ul><li>Three-ring structure with a bridgehead amine, -NH2</li> <li>The -NH2 group is protonated under physiological pH of the body to carry a positive charge, -NH3+</li> <li>Two methyl (CH3) side groups which serve to prolong the dwell time and increase stability as well as affinity for the NMDA receptor channel compared with amantadine (1-adamantanamine).<a href="#cite_note-Lipton1-18">[18]</a><a href="#cite_note-Sonkusare-114">[114]</a></li></ul> Despite the small structural difference between memantine and amantadine, two adamantane derivatives, the affinity for the binding site of NR1/NR2B subunit is much greater for memantine. In <a href="/wiki/Patch-clamp" class="mw-redirect" title="Patch-clamp">patch-clamp</a> measurements memantine has an <a href="/wiki/IC50" title="IC50">IC50</a> of (2.3+0.3) μM while amantadine has an IC50 of (71.0+11.1) μM.<a href="#cite_note-Wanka-28">[28]</a> The binding site with the highest affinity is called the dominant binding site. It involves a connection between the amine group of memantine and the NR1-N161 binding pocket of the NR1/NR2B subunit. The methyl side groups play an important role in increasing the affinity to the open NMDA receptor channels and making it a much better neuroprotective drug than amantadine. The binding pockets for the methyl groups are considered to be at the NR1-A645 and NR2B-A644 of the NR1/NR2B.<a href="#cite_note-Limapichat-32">[32]</a> The binding pockets are shown in figure 2. Memantine binds at or near to the Mg2+ site inside the NMDA receptor associated channel. The -NH2 group on memantine, which is protonated under physiological pH of the body, represents the region that binds at or near to the Mg2+ site.<a href="#cite_note-Lipton1-18">[18]</a> Adding two methyl groups to the -N on the memantine structure has shown to decrease affinity, giving an IC50 value of (28.4+1.4) μM.<a href="#cite_note-Wanka-28">[28]</a> <div class="mw-heading mw-heading4"><h4 id="Second_generation_derivative_of_memantine;_nitromemantine">Second generation derivative of memantine; nitromemantine</h4>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=31" title="Edit section: Second generation derivative of memantine; nitromemantine">edit</a>]</div> Several derivatives of Nitromemantine, a second-generation derivative of memantine, have been synthesized in order to perform a detailed <a href="/wiki/Structure_activity_relationship" class="mw-redirect" title="Structure activity relationship">structure activity relationship</a> (SAR) of these novel drugs. One class, containing a nitro (NO2) group opposite to the bridgehead amine (NH2), showed a promising outcome. Nitromemantine utilizes memantine binding site on the NMDA receptor to target the NOx (X= 1 or 2) group for interaction with the S- nitrosylation/redox site external to the memantine binding site. Lengthening the side chains of memantine compensates for the worse drug affinity in the channel associated with the addition of the –ONO2 group<a href="#cite_note-Takahashi-115">[115]</a> <div class="mw-heading mw-heading3"><h3 id="Therapeutic_application">Therapeutic application</h3>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=32" title="Edit section: Therapeutic application">edit</a>]</div> Excitotoxicity is implied to be involved in some neurodegenerative disorders such as Alzheimer's disease, Parkinson's disease, Huntington's disease and <a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">amyotrophic lateral sclerosis</a>.<a href="#cite_note-Chen-16">[16]</a><a href="#cite_note-Kemp-17">[17]</a><a href="#cite_note-Lipton1-18">[18]</a><a href="#cite_note-Koch-19">[19]</a> Blocking of NMDA receptors could therefore, in theory, be useful in treating such diseases.<a href="#cite_note-Chen-16">[16]</a><a href="#cite_note-Kemp-17">[17]</a><a href="#cite_note-Lipton1-18">[18]</a> It is, however, important to preserve physiological NMDA receptor activity while trying to block its excessive, excitotoxic activity. This can possibly be achieved by uncompetitive antagonists, blocking the receptor's ion channel when excessively open <a href="#cite_note-Lipton1-18">[18]</a> Memantine is an example of uncompetitive NMDA receptor antagonist that has approved indication for the neurodegenerative disease Alzheimer's disease. In 2015 memantine is still in clinical trials for additional neurological diseases.<a href="#cite_note-Limapichat-32">[32]</a><a href="#cite_note-Lipton3-98">[98]</a> <div class="mw-heading mw-heading2"><h2 id="Receptor_modulation">Receptor modulation</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=33" title="Edit section: Receptor modulation">edit</a>]</div> The NMDA receptor is a non-specific cation channel that can allow the passage of Ca2+ and Na+ into the cell and K+ out of the cell. The <a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">excitatory postsynaptic potential</a> (EPSP) produced by activation of an NMDA receptor increases the concentration of Ca2+ in the cell. The Ca2+ can in turn function as a <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">second messenger</a> in various <a href="/wiki/Signaling_pathway" class="mw-redirect" title="Signaling pathway">signaling pathways</a>. However, the NMDA receptor cation channel is blocked by Mg2+ at resting membrane potential.<a href="#cite_note-Purves129-131-116">[116]</a> Magnesium unblock is not instantaneous; to unblock all available channels, the postsynaptic cell must be depolarized for a sufficiently long period of time (in the scale of milliseconds).<a href="#cite_note-Vargas-Caballero-117">[117]</a> Therefore, the NMDA receptor functions as a "molecular <a href="/wiki/Coincidence_detection_in_neurobiology" title="Coincidence detection in neurobiology">coincidence detector</a>". Its ion channel opens only when the following two conditions are met: glutamate is bound to the receptor, and the postsynaptic cell is depolarized (which removes the Mg2+ blocking the channel). This property of the NMDA receptor explains many aspects of <a href="/wiki/Long-term_potentiation" title="Long-term potentiation">long-term potentiation</a> (LTP) and <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a>.<a href="#cite_note-Purves191-195-118">[118]</a> In a <a href="/wiki/Resting_membrane_potential" class="mw-redirect" title="Resting membrane potential">resting-membrane potential</a>, the NMDA receptor pore is opened allowing for an influx of external magnesium ions binding to prevent further ion permeation.<a href="#cite_note-119">[119]</a> External magnesium ions are in a <a href="/wiki/Millimolar" class="mw-redirect" title="Millimolar">millimolar</a> range while intracellular magnesium ions are at a <a href="/wiki/Micromolar" class="mw-redirect" title="Micromolar">micromolar</a> range to result in negative membrane potential. NMDA receptors are modulated by a number of <a href="/wiki/Endogenous" class="mw-redirect" title="Endogenous">endogenous</a> and <a href="/wiki/Exogenous" class="mw-redirect" title="Exogenous">exogenous</a> compounds and play a key role in a wide range of <a href="/wiki/Physiology" title="Physiology">physiological</a> (e.g., <a href="/wiki/Memory" title="Memory">memory</a>) and <a href="/wiki/Pathology" title="Pathology">pathological</a> processes (e.g., <a href="/wiki/Excitotoxicity" title="Excitotoxicity">excitotoxicity</a>). Magnesium works to potentiate NMDA-induced responses at positive membrane potentials while blocking the NMDA channel. The use of calcium, potassium, and sodium are used to modulate the activity of NMDARs passing through the NMDA membrane. Changes in H+ concentration can partially inhibit the activity of NMDA receptors in different physiological conditions. <div class="mw-heading mw-heading2"><h2 id="Clinical_significance">Clinical significance</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=34" title="Edit section: Clinical significance">edit</a>]</div> NMDAR antagonists like <a href="/wiki/Ketamine" title="Ketamine">ketamine</a>, <a href="/wiki/Esketamine" title="Esketamine">esketamine</a>, <a href="/wiki/Tiletamine" title="Tiletamine">tiletamine</a>, <a href="/wiki/Phencyclidine" title="Phencyclidine">phencyclidine</a>, <a href="/wiki/Nitrous_oxide" title="Nitrous oxide">nitrous oxide</a>, and <a href="/wiki/Xenon" title="Xenon">xenon</a> are used as <a href="/wiki/General_anesthetic" class="mw-redirect" title="General anesthetic">general anesthetics</a>. These and similar drugs like <a href="/wiki/Dextromethorphan" title="Dextromethorphan">dextromethorphan</a> and <a href="/wiki/Methoxetamine" title="Methoxetamine">methoxetamine</a> also produce <a href="/wiki/Dissociative_drug" class="mw-redirect" title="Dissociative drug">dissociative</a>, <a href="/wiki/Hallucinogen" title="Hallucinogen">hallucinogenic</a>, and <a href="/wiki/Euphoriant" class="mw-redirect" title="Euphoriant">euphoriant</a> effects and are used as <a href="/wiki/Recreational_drug" class="mw-redirect" title="Recreational drug">recreational drugs</a>. NMDAR-targeted compounds, including ketamine, <a href="/wiki/Esketamine" title="Esketamine">esketamine</a> (JNJ-54135419), <a href="/wiki/Rapastinel" title="Rapastinel">rapastinel</a> (GLYX-13), <a href="/wiki/Apimostinel" title="Apimostinel">apimostinel</a> (NRX-1074), <a href="/wiki/AGN-241751" class="mw-redirect" title="AGN-241751">zelquistinel</a> (AGN-241751), <a href="/wiki/4-chlorokynurenine" class="mw-redirect" title="4-chlorokynurenine">4-chlorokynurenine</a> (AV-101), and <a href="/wiki/Rislenemdaz" title="Rislenemdaz">rislenemdaz</a> (CERC-301, MK-0657), are under development for the treatment of <a href="/wiki/Mood_disorder" title="Mood disorder">mood disorders</a>, including <a href="/wiki/Major_depressive_disorder" title="Major depressive disorder">major depressive disorder</a> and <a href="/wiki/Treatment-resistant_depression" title="Treatment-resistant depression">treatment-resistant depression</a>.<a href="#cite_note-Flight2013-83">[83]</a><a href="#cite_note-VécseiSzalárdy2012-84">[84]</a><a href="#cite_note-issn2168-9709-120">[120]</a> In addition, ketamine is already employed for this purpose as an off-label therapy in some clinics.<a href="#cite_note-NPR2014-121">[121]</a><a href="#cite_note-ScientificAmerican2013-122">[122]</a> Research suggests that <a href="/wiki/Tianeptine" title="Tianeptine">tianeptine</a> produces antidepressant effects through indirect alteration and inhibition of <a href="/wiki/Glutamate_(neurotransmitter)" title="Glutamate (neurotransmitter)">glutamate</a> receptor activity and release of <a href="/wiki/Brain-derived_neurotrophic_factor" title="Brain-derived neurotrophic factor"><abbr title="brain-derived neurotrophic factor">BDNF</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip brain-derived neurotrophic factor, in turn affecting <a href="/wiki/Neural_plasticity" class="mw-redirect" title="Neural plasticity">neural plasticity</a>.<a href="#cite_note-mp092-123">[123]</a><a href="#cite_note-pmid15550348-124">[124]</a><a href="#cite_note-pmid15753957-125">[125]</a><a href="#cite_note-pmid18221189-126">[126]</a><a href="#cite_note-CNS20082-127">[127]</a> Tianeptine also acts on the NMDA and <a href="/wiki/AMPA_receptor" title="AMPA receptor">AMPA receptors</a>.<a href="#cite_note-mp092-123">[123]</a><a href="#cite_note-CNS20082-127">[127]</a> In animal models, tianeptine inhibits the pathological stress-induced changes in glutamatergic neurotransmission in the amygdala and hippocampus. <a href="/wiki/Memantine" title="Memantine">Memantine</a>, a low-trapping NMDAR antagonist, is approved in the <a href="/wiki/United_States" title="United States">United States</a> and <a href="/wiki/Europe" title="Europe">Europe</a> for the treatment of moderate-to-severe Alzheimer's disease,<a href="#cite_note-MountC2006-128">[128]</a> and has now received a limited recommendation by the UK's <a href="/wiki/National_Institute_for_Health_and_Care_Excellence" title="National Institute for Health and Care Excellence">National Institute for Health and Care Excellence</a> for patients who fail other treatment options.<a href="#cite_note-NICE_Guidelines-129">[129]</a> Cochlear NMDARs are the target of intense research to find pharmacological solutions to treat <a href="/wiki/Tinnitus" title="Tinnitus">tinnitus</a>. NMDARs are associated with a rare <a href="/wiki/Autoimmune" class="mw-redirect" title="Autoimmune">autoimmune</a> disease, <a href="/wiki/Anti-NMDA_receptor_encephalitis" title="Anti-NMDA receptor encephalitis">anti-NMDA receptor encephalitis</a> (also known as NMDAR encephalitis<a href="#cite_note-130">[130]</a>), that usually occurs due to cross-reactivity of antibodies produced by the immune system against ectopic brain tissues, such as those found in <a href="/wiki/Teratoma" title="Teratoma">teratoma</a>. These are known as <a href="/wiki/Anti-glutamate_receptor_antibodies" title="Anti-glutamate receptor antibodies">anti-glutamate receptor antibodies</a>. Compared to <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> <a href="/wiki/Stimulant" title="Stimulant">stimulants</a> like <a href="/wiki/Methamphetamine" title="Methamphetamine">methamphetamine</a>, the NMDAR antagonist phencyclidine can produce a wider range of symptoms that resemble schizophrenia in healthy volunteers, in what has led to the <a href="/wiki/Glutamate_hypothesis_of_schizophrenia" title="Glutamate hypothesis of schizophrenia">glutamate hypothesis of schizophrenia</a>.<a href="#cite_note-pmid18395805-131">[131]</a> Experiments in which rodents are treated with NMDA receptor antagonist are today the most common model when it comes to testing of novel schizophrenia therapies or exploring the exact mechanism of drugs already approved for treatment of schizophrenia. NMDAR antagonists, for instance <a href="/wiki/Eliprodil" title="Eliprodil">eliprodil</a>, <a href="/wiki/Gavestinel" title="Gavestinel">gavestinel</a>, <a href="/wiki/Licostinel" title="Licostinel">licostinel</a>, and <a href="/wiki/Selfotel" title="Selfotel">selfotel</a> have been extensively investigated for the treatment of <a href="/wiki/Excitotoxicity" title="Excitotoxicity">excitotoxicity</a>-mediated <a href="/wiki/Neurotoxicity" title="Neurotoxicity">neurotoxicity</a> in situations like <a href="/wiki/Ischemic_stroke" class="mw-redirect" title="Ischemic stroke">ischemic stroke</a> and <a href="/wiki/Traumatic_brain_injury" title="Traumatic brain injury">traumatic brain injury</a>, but were unsuccessful in <a href="/wiki/Clinical_trial" title="Clinical trial">clinical trials</a> used in small doses to avoid sedation, but NMDAR antagonists can block <a href="/wiki/Cortical_spreading_depression" title="Cortical spreading depression">Spreading Depolarizations</a> in animals and in patients with brain injury.<a href="#cite_note-132">[132]</a> This use has not been tested in clinical trials yet. <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=35" title="Edit section: See also">edit</a>]</div> <ul><li><a href="/wiki/Ca2%2B/calmodulin-dependent_protein_kinase" class="mw-redirect" title="Ca2+/calmodulin-dependent protein kinase">Calcium/calmodulin-dependent protein kinases</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=NMDA_receptor&action=edit&section=36" title="Edit section: References">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Laube-1">^ <a href="#cite_ref-Laube_1-0">a</a> <a href="#cite_ref-Laube_1-1">b</a> <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 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AntiqueWhite"><a href="/wiki/Cys-loop_receptors" class="mw-redirect" title="Cys-loop receptors">Cys-loop receptors</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;"><a href="/wiki/5-HT_receptor" title="5-HT receptor">5-HT/serotonin</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/5-HT3_receptor" title="5-HT3 receptor">5-HT3</a> <ul><li><a href="/wiki/HTR3A" title="HTR3A">A</a></li> <li><a href="/wiki/HTR3B" title="HTR3B">B</a></li> <li><a href="/wiki/HTR3C" title="HTR3C">C</a></li> <li><a href="/wiki/HTR3D" title="HTR3D">D</a></li> <li><a href="/wiki/HTR3E" title="HTR3E">E</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;"><a href="/wiki/GABA_receptor" title="GABA receptor">GABA</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/GABAA_receptor" title="GABAA receptor">GABAA</a> <ul><li><a href="/wiki/Gamma-aminobutyric_acid_receptor_subunit_alpha-1" title="Gamma-aminobutyric acid receptor subunit alpha-1">α1</a></li> <li><a href="/wiki/GABRA2" title="GABRA2">α2</a></li> <li><a href="/wiki/GABRA3" title="GABRA3">α3</a></li> <li><a href="/wiki/GABRA4" title="GABRA4">α4</a></li> <li><a href="/wiki/GABRA5" title="GABRA5">α5</a></li> <li><a href="/wiki/GABRA6" title="GABRA6">α6</a></li> <li><a href="/wiki/GABRB1" title="GABRB1">β1</a></li> <li><a href="/wiki/GABRB2" title="GABRB2">β2</a></li> <li><a href="/wiki/GABRB3" title="GABRB3">β3</a></li> <li><a href="/wiki/GABRG1" title="GABRG1">γ1</a></li> <li><a href="/wiki/GABRG2" class="mw-redirect" title="GABRG2">γ2</a></li> <li><a href="/wiki/GABRG3" title="GABRG3">γ3</a></li> <li><a href="/wiki/GABRD" title="GABRD">δ</a></li> <li><a href="/wiki/GABRE" title="GABRE">ε</a></li> <li><a href="/wiki/GABRP" title="GABRP">π</a></li> <li><a href="/wiki/GABRQ" title="GABRQ">θ</a></li></ul></li> <li><a href="/wiki/GABAA-rho_receptor" title="GABAA-rho receptor">GABAA-ρ</a> <ul><li><a href="/wiki/GABRR1" title="GABRR1">ρ1</a></li> <li><a href="/wiki/GABRR2" title="GABRR2">ρ2</a></li> <li><a href="/wiki/GABRR3" title="GABRR3">ρ3</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;"><a href="/wiki/Glycine_receptor" title="Glycine receptor">Glycine</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/Glycine_receptor,_alpha_1" title="Glycine receptor, alpha 1">α1</a></li> <li><a href="/wiki/GLRA2" title="GLRA2">α2</a></li> <li><a href="/wiki/GLRA3" title="GLRA3">α3</a></li> <li><a href="/wiki/GLRA4" title="GLRA4">α4</a></li> <li><a href="/wiki/GLRB" title="GLRB">β</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;"><a href="/wiki/Nicotinic_acetylcholine_receptor" title="Nicotinic acetylcholine receptor">Nicotinic acetylcholine</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li>monomers: <a href="/wiki/CHRNA1" title="CHRNA1">α1</a></li> <li><a href="/wiki/CHRNA2" title="CHRNA2">α2</a></li> <li><a href="/wiki/CHRNA3" title="CHRNA3">α3</a></li> <li><a href="/wiki/CHRNA4" title="CHRNA4">α4</a></li> <li><a href="/wiki/CHRNA5" class="mw-redirect" title="CHRNA5">α5</a></li> <li><a href="/wiki/CHRNA6" title="CHRNA6">α6</a></li> <li><a href="/wiki/CHRNA7" title="CHRNA7">α7</a></li> <li><a href="/wiki/CHRNA9" title="CHRNA9">α9</a></li> <li><a href="/wiki/CHRNA10" title="CHRNA10">α10</a></li> <li><a href="/wiki/CHRNB1" title="CHRNB1">β1</a></li> <li><a href="/wiki/CHRNB2" title="CHRNB2">β2</a></li> <li><a href="/wiki/CHRNB3" title="CHRNB3">β3</a></li> <li><a href="/wiki/CHRNB4" title="CHRNB4">β4</a></li> <li><a href="/wiki/CHRND" title="CHRND">δ</a></li> <li><a href="/wiki/CHRNE" title="CHRNE">ε</a></li></ul> <ul><li>pentamers: <a href="/wiki/Alpha-3_beta-4_nicotinic_receptor" title="Alpha-3 beta-4 nicotinic receptor">(α3)2(β4)3</a></li> <li><a href="/wiki/Alpha-4_beta-2_nicotinic_receptor" title="Alpha-4 beta-2 nicotinic receptor">(α4)2(β2)3</a></li> <li><a href="/wiki/Alpha-7_nicotinic_receptor" title="Alpha-7 nicotinic receptor">(α7)5</a></li> <li><a href="/wiki/Ganglion_type_nicotinic_receptor" class="mw-redirect" title="Ganglion type nicotinic receptor">(α1)2(β4)3 - Ganglion type</a></li> <li><a href="/wiki/Muscle_type_nicotinic_receptor" class="mw-redirect" title="Muscle type nicotinic receptor">(α1)2β1δε - Muscle type</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;">Zinc</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/Zinc-activated_ion_channel" title="Zinc-activated ion channel">Zinc-activated</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">Ionotropic</a> <a href="/wiki/Glutamate_receptor" title="Glutamate receptor">glutamates</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;">Ligand-gated only</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/AMPA_receptor" title="AMPA receptor">AMPA</a> (<a href="/wiki/GRIA1" title="GRIA1">1</a></li> <li><a href="/wiki/GRIA2" title="GRIA2">2</a></li> <li><a href="/wiki/GRIA3" title="GRIA3">3</a></li> <li><a href="/wiki/GRIA4" title="GRIA4">4</a>)</li> <li><a href="/wiki/Kainate_receptor" title="Kainate receptor">Kainate</a> <ul><li><a href="/wiki/GRIK1" title="GRIK1">1</a></li> <li><a href="/wiki/GRIK2" title="GRIK2">2</a></li> <li><a href="/wiki/GRIK3" title="GRIK3">3</a></li> <li><a href="/wiki/GRIK4" title="GRIK4">4</a></li> <li><a href="/wiki/GRIK5" title="GRIK5">5</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;">Voltage- and ligand-gated</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">NMDA</a> <ul><li><a href="/wiki/GRIN1" title="GRIN1">1</a></li> <li><a href="/wiki/GRIN2A" title="GRIN2A">2A</a></li> <li><a href="/wiki/GRIN2B" title="GRIN2B">2B</a></li> <li><a href="/wiki/GRIN2C" title="GRIN2C">2C</a></li> <li><a href="/wiki/GRIN2D" title="GRIN2D">2D</a></li> <li><a href="/wiki/GRIN3A" title="GRIN3A">3A</a></li> <li><a href="/wiki/GRIN3B" title="GRIN3B">3B</a></li> <li><a href="/wiki/GRINL1A" title="GRINL1A">L1A</a></li> <li><a href="/wiki/GRINL1B" title="GRINL1B">L1B</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;">‘Orphan’</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li>GluD <ul><li><a href="/wiki/GRID1" title="GRID1">δ1</a></li> <li><a href="/wiki/GRID2" title="GRID2">δ2</a></li></ul></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/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a>-gated channels</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Purinergic_receptors" scope="row" class="navbox-group" style="width:14em;background-color: AntiqueWhite;"><a href="/wiki/Purinergic_receptor" title="Purinergic receptor">Purinergic receptors</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/P2X_receptor" class="mw-redirect" title="P2X receptor">P2X</a> <ul><li><a href="/wiki/P2RX1" title="P2RX1">1</a></li> <li><a href="/wiki/P2RX2" title="P2RX2">2</a></li> <li><a href="/wiki/P2RX3" title="P2RX3">3</a></li> <li><a href="/wiki/P2RX4" title="P2RX4">4</a></li> <li><a href="/wiki/P2RX5" title="P2RX5">5</a></li> <li><a href="/wiki/P2RX6" title="P2RX6">6</a></li> <li><a href="/wiki/P2RX7" title="P2RX7">7</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ionotropic_glutamate_receptor_modulators" style="padding:3px"><table class="nowraplinks hlist 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:Ionotropic_glutamate_receptor_modulators" title="Template:Ionotropic glutamate receptor modulators"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Ionotropic_glutamate_receptor_modulators" title="Template talk:Ionotropic glutamate receptor modulators"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Ionotropic_glutamate_receptor_modulators" title="Special:EditPage/Template:Ionotropic glutamate receptor modulators"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ionotropic_glutamate_receptor_modulators" style="font-size:114%;margin:0 4em"><a href="/wiki/Ionotropic_glutamate_receptor" title="Ionotropic glutamate receptor">Ionotropic glutamate receptor</a> <a href="/wiki/Receptor_modulator" title="Receptor modulator">modulators</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/%CE%91-Amino-3-hydroxy-5-methyl-4-isoxazolepropionic_acid_receptor" class="mw-redirect" title="Α-Amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor"><abbr title="α-Amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor">AMPAR</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip α-Amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Agonists: Main site agonists: <a href="/wiki/5-Fluorowillardiine" title="5-Fluorowillardiine">5-Fluorowillardiine</a></li> <li><a href="/wiki/Acromelic_acid_A" title="Acromelic acid A">Acromelic acid (acromelate)</a></li> <li><a href="/wiki/AMPA" title="AMPA">AMPA</a></li> <li><a href="/w/index.php?title=%CE%92-N-Oxalylamino-L-Alanine&action=edit&redlink=1" class="new" title="Β-N-Oxalylamino-L-Alanine (page does not exist)">BOAA</a></li> <li><a href="/wiki/Domoic_acid" title="Domoic acid">Domoic acid</a></li> <li><a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">Glutamate</a></li> <li><a href="/wiki/Ibotenic_acid" title="Ibotenic acid">Ibotenic acid</a></li> <li><a href="/wiki/Proline" title="Proline">Proline</a></li> <li><a href="/wiki/Quisqualic_acid" title="Quisqualic acid">Quisqualic acid</a></li> <li><a href="/wiki/Willardiine" title="Willardiine">Willardiine</a>; Positive allosteric modulators: <a href="/wiki/Aniracetam" title="Aniracetam">Aniracetam</a></li> <li><a href="/wiki/Cyclothiazide" title="Cyclothiazide">Cyclothiazide</a></li> <li><a href="/wiki/CX-516" title="CX-516">CX-516</a></li> <li><a href="/wiki/CX-546" title="CX-546">CX-546</a></li> <li><a href="/wiki/CX614" title="CX614">CX-614</a></li> <li><a href="/wiki/Farampator" title="Farampator">Farampator (CX-691, ORG-24448)</a></li> <li><a href="/wiki/CX717" title="CX717">CX-717</a></li> <li><a href="/wiki/CX-1739" class="mw-redirect" title="CX-1739">CX-1739</a></li> <li><a href="/w/index.php?title=CX-1942&action=edit&redlink=1" class="new" title="CX-1942 (page does not exist)">CX-1942</a></li> <li><a href="/wiki/Diazoxide" title="Diazoxide">Diazoxide</a></li> <li><a href="/wiki/Hydrochlorothiazide" title="Hydrochlorothiazide">Hydrochlorothiazide (HCTZ)</a></li> <li><a href="/wiki/IDRA-21" title="IDRA-21">IDRA-21</a></li> <li><a href="/w/index.php?title=LY-392098&action=edit&redlink=1" class="new" title="LY-392098 (page does not exist)">LY-392098</a></li> <li><a href="/w/index.php?title=LY-395153&action=edit&redlink=1" class="new" title="LY-395153 (page does not exist)">LY-395153</a></li> <li><a href="/wiki/LY-404187" title="LY-404187">LY-404187</a></li> <li><a href="/w/index.php?title=LY-451646&action=edit&redlink=1" class="new" title="LY-451646 (page does not exist)">LY-451646</a></li> <li><a href="/wiki/LY-503430" title="LY-503430">LY-503430</a></li> <li><a href="/wiki/Mibampator" title="Mibampator">Mibampator (LY-451395)</a></li> <li><a href="/wiki/Nooglutyl" title="Nooglutyl">Nooglutyl</a></li> <li><a href="/wiki/ORG-26576" title="ORG-26576">ORG-26576</a></li> <li><a href="/wiki/Oxiracetam" title="Oxiracetam">Oxiracetam</a></li> <li><a href="/wiki/PEPA_(drug)" title="PEPA (drug)">PEPA</a></li> <li><a href="/wiki/Pesampator" title="Pesampator">Pesampator (BIIB-104, PF-04958242)</a></li> <li><a href="/wiki/Piracetam" title="Piracetam">Piracetam</a></li> <li><a href="/wiki/Pramiracetam" title="Pramiracetam">Pramiracetam</a></li> <li><a href="/wiki/S-18986" title="S-18986">S-18986</a></li> <li><a href="/wiki/Tulrampator" title="Tulrampator">Tulrampator (S-47445, CX-1632)</a></li></ul> <ul><li>Antagonists: <a href="/w/index.php?title=ACEA-1011&action=edit&redlink=1" class="new" title="ACEA-1011 (page does not exist)">ACEA-1011</a></li> <li><a href="/w/index.php?title=ATPO&action=edit&redlink=1" class="new" title="ATPO (page does not exist)">ATPO</a></li> <li><a href="/wiki/Becampanel" title="Becampanel">Becampanel</a></li> <li><a href="/wiki/Caroverine" title="Caroverine">Caroverine</a></li> <li><a href="/wiki/CNQX" title="CNQX">CNQX</a></li> <li><a href="/wiki/Dasolampanel" title="Dasolampanel">Dasolampanel</a></li> <li><a href="/wiki/DNQX" title="DNQX">DNQX</a></li> <li><a href="/wiki/Fanapanel" title="Fanapanel">Fanapanel (MPQX)</a></li> <li><a href="/w/index.php?title=Gamma-D-Glutamylaminomethylsulfonic_acid&action=edit&redlink=1" class="new" title="Gamma-D-Glutamylaminomethylsulfonic acid (page does not exist)">GAMS</a></li> <li><a href="/wiki/Kaitocephalin" title="Kaitocephalin">Kaitocephalin</a></li> <li><a href="/wiki/Kynurenic_acid" title="Kynurenic acid">Kynurenic acid</a></li> <li><a href="/wiki/Kynurenine" title="Kynurenine">Kynurenine</a></li> <li><a href="/wiki/Licostinel" title="Licostinel">Licostinel (ACEA-1021)</a></li> <li><a href="/wiki/NBQX" title="NBQX">NBQX</a></li> <li><a href="/w/index.php?title=PNQX&action=edit&redlink=1" class="new" title="PNQX (page does not exist)">PNQX</a></li> <li><a href="/wiki/Selurampanel" title="Selurampanel">Selurampanel</a></li> <li><a href="/wiki/Tezampanel" title="Tezampanel">Tezampanel</a></li> <li><a href="/wiki/Theanine" title="Theanine">Theanine</a></li> <li><a href="/wiki/Topiramate" title="Topiramate">Topiramate</a></li> <li><a href="/w/index.php?title=YM90K&action=edit&redlink=1" class="new" title="YM90K (page does not exist)">YM90K</a></li> <li><a href="/wiki/Zonampanel" title="Zonampanel">Zonampanel</a>; Negative allosteric modulators: <a href="/wiki/Barbiturates" class="mw-redirect" title="Barbiturates">Barbiturates</a> (e.g., <a href="/wiki/Pentobarbital" title="Pentobarbital">pentobarbital</a>, <a href="/wiki/Sodium_thiopental" title="Sodium thiopental">sodium thiopental</a>)</li> <li><a href="/wiki/Cyclopropane" title="Cyclopropane">Cyclopropane</a></li> <li><a href="/wiki/Enflurane" title="Enflurane">Enflurane</a></li> <li><a href="/wiki/Alcohol_(drug)" title="Alcohol (drug)">Ethanol (alcohol)</a></li> <li><a href="/wiki/Evans_blue_(dye)" title="Evans blue (dye)">Evans blue</a></li> <li><a href="/wiki/GYKI_52466" title="GYKI 52466">GYKI-52466</a></li> <li><a href="/w/index.php?title=GYKI-53655&action=edit&redlink=1" class="new" title="GYKI-53655 (page does not exist)">GYKI-53655</a></li> <li><a href="/wiki/Halothane" title="Halothane">Halothane</a></li> <li><a href="/wiki/Irampanel" title="Irampanel">Irampanel</a></li> <li><a href="/wiki/Isoflurane" title="Isoflurane">Isoflurane</a></li> <li><a href="/wiki/Perampanel" title="Perampanel">Perampanel</a></li> <li><a href="/wiki/Pregnenolone_sulfate" title="Pregnenolone sulfate">Pregnenolone sulfate</a></li> <li><a href="/wiki/Sevoflurane" title="Sevoflurane">Sevoflurane</a></li> <li><a href="/wiki/Talampanel" title="Talampanel">Talampanel</a>; Unknown/unsorted antagonists: <a href="/wiki/Minocycline" title="Minocycline">Minocycline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/Kainate_receptor" title="Kainate receptor"><abbr title="Kainate receptor">KAR</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip Kainate receptor</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Agonists: Main site agonists: <a href="/w/index.php?title=5-Bromowillardiine&action=edit&redlink=1" class="new" title="5-Bromowillardiine (page does not exist)">5-Bromowillardiine</a></li> <li><a href="/wiki/5-Iodowillardiine" title="5-Iodowillardiine">5-Iodowillardiine</a></li> <li><a href="/w/index.php?title=Acromelic_acid&action=edit&redlink=1" class="new" title="Acromelic acid (page does not exist)">Acromelic acid (acromelate)</a></li> <li><a href="/wiki/AMPA" title="AMPA">AMPA</a></li> <li><a href="/w/index.php?title=ATPA&action=edit&redlink=1" class="new" title="ATPA (page does not exist)">ATPA</a></li> <li><a href="/wiki/Domoic_acid" title="Domoic acid">Domoic acid</a></li> <li><a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">Glutamate</a></li> <li><a href="/wiki/Ibotenic_acid" title="Ibotenic acid">Ibotenic acid</a></li> <li><a href="/wiki/Kainic_acid" title="Kainic acid">Kainic acid</a></li> <li><a href="/w/index.php?title=LY-339434&action=edit&redlink=1" class="new" title="LY-339434 (page does not exist)">LY-339434</a></li> <li><a href="/wiki/Proline" title="Proline">Proline</a></li> <li><a href="/wiki/Quisqualic_acid" title="Quisqualic acid">Quisqualic acid</a></li> <li><a href="/wiki/SYM-2081" title="SYM-2081">SYM-2081</a>; Positive allosteric modulators: <a href="/wiki/Cyclothiazide" title="Cyclothiazide">Cyclothiazide</a></li> <li><a href="/wiki/Diazoxide" title="Diazoxide">Diazoxide</a></li> <li><a href="/wiki/Enflurane" title="Enflurane">Enflurane</a></li> <li><a href="/wiki/Halothane" title="Halothane">Halothane</a></li> <li><a href="/wiki/Isoflurane" title="Isoflurane">Isoflurane</a></li></ul> <ul><li>Antagonists: <a href="/w/index.php?title=ACEA-1011&action=edit&redlink=1" class="new" title="ACEA-1011 (page does not exist)">ACEA-1011</a></li> <li><a href="/wiki/CNQX" title="CNQX">CNQX</a></li> <li><a href="/wiki/Dasolampanel" title="Dasolampanel">Dasolampanel</a></li> <li><a href="/wiki/DNQX" title="DNQX">DNQX</a></li> <li><a href="/w/index.php?title=Gamma-D-Glutamylaminomethylsulfonic_acid&action=edit&redlink=1" class="new" title="Gamma-D-Glutamylaminomethylsulfonic acid (page does not exist)">GAMS</a></li> <li><a href="/wiki/Kaitocephalin" title="Kaitocephalin">Kaitocephalin</a></li> <li><a href="/wiki/Kynurenic_acid" title="Kynurenic acid">Kynurenic acid</a></li> <li><a href="/wiki/Licostinel" title="Licostinel">Licostinel (ACEA-1021)</a></li> <li><a href="/w/index.php?title=LY-382884&action=edit&redlink=1" class="new" title="LY-382884 (page does not exist)">LY-382884</a></li> <li><a href="/wiki/NBQX" title="NBQX">NBQX</a></li> <li><a href="/wiki/NS102" title="NS102">NS102</a></li> <li><a href="/wiki/Selurampanel" title="Selurampanel">Selurampanel</a></li> <li><a href="/wiki/Tezampanel" title="Tezampanel">Tezampanel</a></li> <li><a href="/wiki/Theanine" title="Theanine">Theanine</a></li> <li><a href="/wiki/Topiramate" title="Topiramate">Topiramate</a></li> <li><a href="/wiki/UBP-302" title="UBP-302">UBP-302</a>; Negative allosteric modulators: <a href="/wiki/Barbiturates" class="mw-redirect" title="Barbiturates">Barbiturates</a> (e.g., <a href="/wiki/Pentobarbital" title="Pentobarbital">pentobarbital</a>, <a href="/wiki/Sodium_thiopental" title="Sodium thiopental">sodium thiopental</a>)</li> <li><a href="/wiki/Enflurane" title="Enflurane">Enflurane</a></li> <li><a href="/wiki/Alcohol_(drug)" title="Alcohol (drug)">Ethanol (alcohol)</a></li> <li><a href="/wiki/Evans_blue_(dye)" title="Evans blue (dye)">Evans blue</a></li> <li><a href="/w/index.php?title=NS-3763&action=edit&redlink=1" class="new" title="NS-3763 (page does not exist)">NS-3763</a></li> <li><a href="/wiki/Pregnenolone_sulfate" title="Pregnenolone sulfate">Pregnenolone sulfate</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/N-Methyl-D-aspartate_receptor" class="mw-redirect" title="N-Methyl-D-aspartate receptor"><abbr title="N-Methyl-D-aspartate receptor">NMDAR</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip N-Methyl-D-aspartate receptor</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Agonists: Main site agonists: <a href="/w/index.php?title=2-Amino-2-(3-hydroxy-5-methylisoxazol-4-yl)acetic_acid&action=edit&redlink=1" class="new" title="2-Amino-2-(3-hydroxy-5-methylisoxazol-4-yl)acetic acid (page does not exist)">AMAA</a></li> <li><a href="/wiki/Aspartate" class="mw-redirect" title="Aspartate">Aspartate</a></li> <li><a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">Glutamate</a></li> <li><a href="/wiki/Homocysteic_acid" title="Homocysteic acid">Homocysteic acid (L-HCA)</a></li> <li><a href="/wiki/Homoquinolinic_acid" title="Homoquinolinic acid">Homoquinolinic acid</a></li> <li><a href="/wiki/Ibotenic_acid" title="Ibotenic acid">Ibotenic acid</a></li> <li><a href="/wiki/N-Methyl-D-aspartic_acid" title="N-Methyl-D-aspartic acid">NMDA</a></li> <li><a href="/wiki/Proline" title="Proline">Proline</a></li> <li><a href="/wiki/Quinolinic_acid" title="Quinolinic acid">Quinolinic acid</a></li> <li><a href="/wiki/Tetrazolylglycine" title="Tetrazolylglycine">Tetrazolylglycine</a></li> <li><a href="/wiki/Theanine" title="Theanine">Theanine</a>; Glycine site agonists: <a href="/w/index.php?title=%CE%92-Fluoro-D-alanine&action=edit&redlink=1" class="new" title="Β-Fluoro-D-alanine (page does not exist)">β-Fluoro-D-alanine</a></li> <li><a href="/w/index.php?title=ACBD_(drug)&action=edit&redlink=1" class="new" title="ACBD (drug) (page does not exist)">ACBD</a></li> <li><a href="/wiki/1-Aminocyclopropanecarboxylic_acid" class="mw-redirect" title="1-Aminocyclopropanecarboxylic acid">ACC (ACPC)</a></li> <li><a href="/wiki/ACPD" title="ACPD">ACPD</a></li> <li><a href="/w/index.php?title=AK-51_(drug)&action=edit&redlink=1" class="new" title="AK-51 (drug) (page does not exist)">AK-51</a></li> <li><a href="/wiki/Apimostinel" title="Apimostinel">Apimostinel (NRX-1074)</a></li> <li><a href="/wiki/B6B21" class="mw-redirect" title="B6B21">B6B21</a></li> <li><a href="/w/index.php?title=Carboxycyclopropylglycine&action=edit&redlink=1" class="new" title="Carboxycyclopropylglycine (page does not exist)">CCG</a></li> <li><a href="/wiki/Alanine" title="Alanine">D-Alanine</a></li> <li><a href="/wiki/Cycloserine" title="Cycloserine">D-Cycloserine</a></li> <li><a href="/wiki/Serine" title="Serine">D-Serine</a></li> <li><a href="/w/index.php?title=N-(3,3-Diphenylpropyl)glycinamide&action=edit&redlink=1" class="new" title="N-(3,3-Diphenylpropyl)glycinamide (page does not exist)">DHPG</a></li> <li><a href="/wiki/Dimethylglycine" title="Dimethylglycine">Dimethylglycine</a></li> <li><a href="/wiki/Glycine" title="Glycine">Glycine</a></li> <li><a href="/wiki/HA-966" title="HA-966">HA-966</a></li> <li><a href="/w/index.php?title=L-687414&action=edit&redlink=1" class="new" title="L-687414 (page does not exist)">L-687414</a></li> <li><a href="/wiki/Alanine" title="Alanine">L-Alanine</a></li> <li><a href="/wiki/Serine" title="Serine">L-Serine</a></li> <li><a href="/wiki/Milacemide" title="Milacemide">Milacemide</a></li> <li><a href="/wiki/Neboglamine" title="Neboglamine">Neboglamine (nebostinel)</a></li> <li><a href="/wiki/Rapastinel" title="Rapastinel">Rapastinel (GLYX-13)</a></li> <li><a href="/wiki/Sarcosine" title="Sarcosine">Sarcosine</a>; Polyamine site agonists: <a href="/wiki/Neomycin" title="Neomycin">Neomycin</a></li> <li><a href="/wiki/Spermidine" title="Spermidine">Spermidine</a></li> <li><a href="/wiki/Spermine" title="Spermine">Spermine</a>; Other positive allosteric modulators: <a href="/wiki/24S-Hydroxycholesterol" title="24S-Hydroxycholesterol">24S-Hydroxycholesterol</a></li> <li><a href="/wiki/Dehydroepiandrosterone" title="Dehydroepiandrosterone"><abbr title="Dehydroepiandrosterone">DHEA</abbr></a><span class="sr-only" style="border: 0; clip: rect(0, 0, 0, 0); clip-path: polygon(0px 0px, 0px 0px, 0px 0px); height: 1px; margin: -1px; overflow: hidden; padding: 0; position: absolute; width: 1px; white-space: nowrap;">Tooltip Dehydroepiandrosterone (<a href="/wiki/Prasterone" title="Prasterone">prasterone</a>)</li> <li><a href="/wiki/Dehydroepiandrosterone_sulfate" title="Dehydroepiandrosterone sulfate"><abbr title="Dehydroepiandrosterone">DHEA</abbr> sulfate</a> (<a href="/wiki/Prasterone_sulfate" title="Prasterone sulfate">prasterone sulfate</a>)</li> <li><a href="/w/index.php?title=Epipregnanolone_sulfate&action=edit&redlink=1" class="new" title="Epipregnanolone sulfate (page does not exist)">Epipregnanolone sulfate</a></li> <li><a href="/w/index.php?title=Plazinemdor&action=edit&redlink=1" class="new" title="Plazinemdor (page does not exist)">Plazinemdor</a></li> <li><a href="/wiki/Pregnenolone_sulfate" title="Pregnenolone sulfate">Pregnenolone sulfate</a></li> <li><a href="/w/index.php?title=SAGE-201&action=edit&redlink=1" class="new" title="SAGE-201 (page does not exist)">SAGE-201</a></li> <li><a href="/w/index.php?title=SAGE-301&action=edit&redlink=1" class="new" title="SAGE-301 (page does not exist)">SAGE-301</a></li> <li><a href="/wiki/SAGE-718" class="mw-redirect" title="SAGE-718">SAGE-718</a></li></ul> <ul><li>Antagonists: Competitive antagonists: <a href="/wiki/AP5" title="AP5">AP5 (APV)</a></li> <li><a href="/wiki/AP-7_(drug)" title="AP-7 (drug)">AP7</a></li> <li><a href="/wiki/CGP-37849" title="CGP-37849">CGP-37849</a></li> <li><a href="/wiki/CGP-39551" title="CGP-39551">CGP-39551</a></li> <li><a href="/w/index.php?title=CGP-39653&action=edit&redlink=1" class="new" title="CGP-39653 (page does not exist)">CGP-39653</a></li> <li><a href="/w/index.php?title=CGP-40116&action=edit&redlink=1" class="new" title="CGP-40116 (page does not exist)">CGP-40116</a></li> <li><a href="/wiki/CGS-19755" class="mw-redirect" title="CGS-19755">CGS-19755</a></li> <li><a href="/w/index.php?title=3-(2-carboxypiperazin-4-yl)propyl-1-phosphonic_acid&action=edit&redlink=1" class="new" title="3-(2-carboxypiperazin-4-yl)propyl-1-phosphonic acid (page does not exist)">CPP</a></li> <li><a href="/wiki/Kaitocephalin" title="Kaitocephalin">Kaitocephalin</a></li> <li><a href="/w/index.php?title=LY-233053&action=edit&redlink=1" class="new" title="LY-233053 (page does not exist)">LY-233053</a></li> <li><a href="/wiki/LY-235959" title="LY-235959">LY-235959</a></li> <li><a href="/w/index.php?title=LY-274614&action=edit&redlink=1" class="new" title="LY-274614 (page does not exist)">LY-274614</a></li> <li><a href="/w/index.php?title=MDL-100453&action=edit&redlink=1" class="new" title="MDL-100453 (page does not exist)">MDL-100453</a></li> <li><a href="/wiki/Midafotel" title="Midafotel">Midafotel (d-CPPene)</a></li> <li><a href="/w/index.php?title=NPC-12626&action=edit&redlink=1" class="new" title="NPC-12626 (page does not exist)">NPC-12626</a></li> <li><a href="/w/index.php?title=NPC-17742&action=edit&redlink=1" class="new" title="NPC-17742 (page does not exist)">NPC-17742</a></li> <li><a href="/w/index.php?title=PBPD&action=edit&redlink=1" class="new" title="PBPD (page does not exist)">PBPD</a></li> <li><a href="/wiki/PEAQX" title="PEAQX">PEAQX</a></li> <li><a href="/wiki/Perzinfotel" title="Perzinfotel">Perzinfotel</a></li> <li><a href="/w/index.php?title=Cis-1-(phenanthrene-2-carbonyl)piperazine-2,3-dicarboxylic_acid&action=edit&redlink=1" class="new" title="Cis-1-(phenanthrene-2-carbonyl)piperazine-2,3-dicarboxylic acid (page does not exist)">PPDA</a></li> <li><a href="/w/index.php?title=SDZ-220581&action=edit&redlink=1" class="new" title="SDZ-220581 (page does not exist)">SDZ-220581</a></li> <li><a href="/wiki/Selfotel" title="Selfotel">Selfotel</a>; Glycine site antagonists: <a href="/wiki/4-Chlorokynurenine" title="4-Chlorokynurenine">4-Cl-KYN (AV-101)</a></li> <li><a href="/wiki/5,7-Dichlorokynurenic_acid" title="5,7-Dichlorokynurenic acid">5,7-DCKA</a></li> <li><a href="/wiki/7-Chlorokynurenic_acid" title="7-Chlorokynurenic acid">7-CKA</a></li> <li><a href="/wiki/1-Aminocyclopropane-1-carboxylic_acid" title="1-Aminocyclopropane-1-carboxylic acid">ACC</a></li> <li><a href="/w/index.php?title=ACEA-1011&action=edit&redlink=1" class="new" title="ACEA-1011 (page does not exist)">ACEA-1011</a></li> <li><a href="/w/index.php?title=ACEA-1328&action=edit&redlink=1" class="new" title="ACEA-1328 (page does not exist)">ACEA-1328</a></li> <li><a href="/wiki/Apimostinel" title="Apimostinel">Apimostinel (NRX-1074)</a></li> <li><a href="/wiki/AV-101" class="mw-redirect" title="AV-101">AV-101</a></li> <li><a href="/wiki/Carisoprodol" title="Carisoprodol">Carisoprodol</a></li> <li><a href="/w/index.php?title=CGP-39653&action=edit&redlink=1" class="new" title="CGP-39653 (page does not exist)">CGP-39653</a></li> <li><a href="/wiki/CNQX" title="CNQX">CNQX</a></li> <li><a href="/wiki/Cycloserine" title="Cycloserine">D-Cycloserine</a></li> <li><a href="/wiki/DNQX" title="DNQX">DNQX</a></li> <li><a href="/wiki/Felbamate" title="Felbamate">Felbamate</a></li> <li><a href="/wiki/Gavestinel" title="Gavestinel">Gavestinel</a></li> <li><a href="/w/index.php?title=GV-196771&action=edit&redlink=1" class="new" title="GV-196771 (page does not exist)">GV-196771</a></li> <li><a href="/w/index.php?title=Harkoseride&action=edit&redlink=1" class="new" title="Harkoseride (page does not exist)">Harkoseride</a></li> <li><a href="/wiki/Kynurenic_acid" title="Kynurenic acid">Kynurenic acid</a></li> <li><a href="/wiki/Kynurenine" title="Kynurenine">Kynurenine</a></li> <li><a href="/w/index.php?title=L-689560&action=edit&redlink=1" class="new" title="L-689560 (page does not exist)">L-689560</a></li> <li><a href="/wiki/L-701324" title="L-701324">L-701324</a></li> <li><a href="/wiki/Licostinel" title="Licostinel">Licostinel (ACEA-1021)</a></li> <li><a href="/w/index.php?title=LU-73068&action=edit&redlink=1" class="new" title="LU-73068 (page does not exist)">LU-73068</a></li> <li><a href="/w/index.php?title=MDL-105519&action=edit&redlink=1" class="new" title="MDL-105519 (page does not exist)">MDL-105519</a></li> <li><a href="/wiki/Meprobamate" title="Meprobamate">Meprobamate</a></li> <li><a href="/w/index.php?title=MRZ_2/576&action=edit&redlink=1" class="new" title="MRZ 2/576 (page does not exist)">MRZ 2/576</a></li> <li><a href="/w/index.php?title=PNQX&action=edit&redlink=1" class="new" title="PNQX (page does not exist)">PNQX</a></li> <li><a href="/wiki/Rapastinel" title="Rapastinel">Rapastinel (GLYX-13)</a></li> <li><a href="/wiki/ZD-9379" title="ZD-9379">ZD-9379</a>; Polyamine site antagonists: <a href="/w/index.php?title=Arcaine&action=edit&redlink=1" class="new" title="Arcaine (page does not exist)">Arcaine</a></li> <li><a href="/w/index.php?title=Co_101676&action=edit&redlink=1" class="new" title="Co 101676 (page does not exist)">Co 101676</a></li> <li><a href="/wiki/1,3-Diaminopropane" title="1,3-Diaminopropane">Diaminopropane</a></li> <li><a href="/wiki/Diethylenetriamine" title="Diethylenetriamine">Diethylenetriamine</a></li> <li><a href="/wiki/Huperzine_A" title="Huperzine A">Huperzine A</a></li> <li><a href="/wiki/Putrescine" title="Putrescine">Putrescine</a>; Uncompetitive pore blockers (mostly dizocilpine site): <a href="/wiki/2-MDP" title="2-MDP">2-MDP</a></li> <li><a href="/wiki/3-HO-PCP" title="3-HO-PCP">3-HO-PCP</a></li> <li><a href="/wiki/3-MeO-PCE" title="3-MeO-PCE">3-MeO-PCE</a></li> <li><a href="/wiki/3-MeO-PCMo" title="3-MeO-PCMo">3-MeO-PCMo</a></li> <li><a href="/wiki/3-MeO-PCP" title="3-MeO-PCP">3-MeO-PCP</a></li> <li><a href="/wiki/4-MeO-PCP" title="4-MeO-PCP">4-MeO-PCP</a></li> <li><a href="/wiki/8a-Phenyldecahydroquinoline" class="mw-redirect" title="8a-Phenyldecahydroquinoline">8A-PDHQ</a></li> <li><a href="/wiki/18-Methoxycoronaridine" title="18-Methoxycoronaridine">18-MC</a></li> <li><a href="/wiki/Alpha-Endopsychosin" class="mw-redirect" title="Alpha-Endopsychosin">α-Endopsychosin</a></li> <li><a href="/wiki/Alaproclate" title="Alaproclate">Alaproclate</a></li> <li><a href="/wiki/Alazocine" title="Alazocine">Alazocine (SKF-10047)</a></li> <li><a href="/wiki/Amantadine" title="Amantadine">Amantadine</a></li> <li><a href="/wiki/Aptiganel" title="Aptiganel">Aptiganel</a></li> <li><a href="/w/index.php?title=Argiotoxin-636&action=edit&redlink=1" class="new" title="Argiotoxin-636 (page does not exist)">Argiotoxin-636</a></li> <li><a href="/wiki/Arketamine" title="Arketamine">Arketamine</a></li> <li><a href="/w/index.php?title=ARL-12495&action=edit&redlink=1" class="new" title="ARL-12495 (page does not exist)">ARL-12495</a></li> <li><a href="/wiki/ARL-15896-AR" class="mw-redirect" title="ARL-15896-AR">ARL-15896-AR</a></li> <li><a href="/wiki/ARL-16247" class="mw-redirect" title="ARL-16247">ARL-16247</a></li> <li><a href="/wiki/Budipine" title="Budipine">Budipine</a></li> <li><a href="/wiki/Coronaridine" title="Coronaridine">Coronaridine</a></li> <li><a href="/wiki/Delucemine" title="Delucemine">Delucemine (NPS-1506)</a></li> <li><a href="/wiki/Dexoxadrol" title="Dexoxadrol">Dexoxadrol</a></li> <li><a href="/wiki/Dextrallorphan" title="Dextrallorphan">Dextrallorphan</a></li> <li><a href="/wiki/Dextromethadone" class="mw-redirect" title="Dextromethadone">Dextromethadone</a></li> <li><a href="/wiki/Dextromethorphan" title="Dextromethorphan">Dextromethorphan</a></li> <li><a href="/wiki/Dextrorphan" title="Dextrorphan">Dextrorphan</a></li> <li><a href="/wiki/Dieticyclidine" title="Dieticyclidine">Dieticyclidine</a></li> <li><a href="/wiki/Diphenidine" title="Diphenidine">Diphenidine</a></li> <li><a href="/wiki/Dizocilpine" title="Dizocilpine">Dizocilpine</a></li> <li><a href="/wiki/Ephenidine" title="Ephenidine">Ephenidine</a></li> <li><a href="/wiki/Esketamine" title="Esketamine">Esketamine</a></li> <li><a href="/wiki/Etoxadrol" title="Etoxadrol">Etoxadrol</a></li> <li><a href="/wiki/Eticyclidine" title="Eticyclidine">Eticyclidine</a></li> <li><a href="/wiki/Fluorolintane" title="Fluorolintane">Fluorolintane</a></li> <li><a href="/wiki/Gacyclidine" title="Gacyclidine">Gacyclidine</a></li> <li><a href="/wiki/Ibogaine" title="Ibogaine">Ibogaine</a></li> <li><a href="/wiki/Ibogamine" title="Ibogamine">Ibogamine</a></li> <li><a href="/wiki/Indantadol" title="Indantadol">Indantadol</a></li> <li><a href="/wiki/Ketamine" title="Ketamine">Ketamine</a></li> <li><a href="/wiki/Ketobemidone" title="Ketobemidone">Ketobemidone</a></li> <li><a href="/wiki/Lanicemine" title="Lanicemine">Lanicemine</a></li> <li><a href="/wiki/Levomethadone" title="Levomethadone">Levomethadone</a></li> <li><a href="/wiki/Levomethorphan" title="Levomethorphan">Levomethorphan</a></li> <li><a href="/wiki/Levomilnacipran" title="Levomilnacipran">Levomilnacipran</a></li> <li><a href="/wiki/Levorphanol" title="Levorphanol">Levorphanol</a></li> <li><a href="/wiki/Loperamide" title="Loperamide">Loperamide</a></li> <li><a href="/wiki/Memantine" title="Memantine">Memantine</a></li> <li><a href="/wiki/Methadone" title="Methadone">Methadone</a></li> <li><a href="/wiki/Methorphan" title="Methorphan">Methorphan</a></li> <li><a href="/wiki/Methoxetamine" title="Methoxetamine">Methoxetamine</a></li> <li><a href="/wiki/Methoxphenidine" title="Methoxphenidine">Methoxphenidine</a></li> <li><a href="/wiki/Milnacipran" title="Milnacipran">Milnacipran</a></li> <li><a href="/wiki/Morphanol" class="mw-redirect" title="Morphanol">Morphanol</a></li> <li><a href="/wiki/NEFA_(drug)" title="NEFA (drug)">NEFA</a></li> <li><a href="/wiki/Neramexane" title="Neramexane">Neramexane</a></li> <li><a href="/wiki/Nitromemantine" title="Nitromemantine">Nitromemantine</a></li> <li><a href="/wiki/Noribogaine" title="Noribogaine">Noribogaine</a></li> <li><a href="/wiki/Norketamine" title="Norketamine">Norketamine</a></li> <li><a href="/wiki/Orphenadrine" title="Orphenadrine">Orphenadrine</a></li> <li><a href="/wiki/PCPr" title="PCPr">PCPr</a></li> <li><a href="/wiki/PD-137889" title="PD-137889">PD-137889</a></li> <li><a href="/wiki/Pethidine" title="Pethidine">Pethidine (meperidine)</a></li> <li><a href="/wiki/Phencyclamine" class="mw-redirect" title="Phencyclamine">Phencyclamine</a></li> <li><a href="/wiki/Phencyclidine" title="Phencyclidine">Phencyclidine</a></li> <li><a href="/wiki/Propoxyphene" class="mw-redirect" title="Propoxyphene">Propoxyphene</a></li> <li><a href="/wiki/Remacemide" title="Remacemide">Remacemide</a></li> <li><a href="/wiki/Rhynchophylline" title="Rhynchophylline">Rhynchophylline</a></li> <li><a href="/wiki/Rimantadine" title="Rimantadine">Rimantadine</a></li> <li><a href="/wiki/Rolicyclidine" title="Rolicyclidine">Rolicyclidine</a></li> <li><a href="/wiki/Sabeluzole" title="Sabeluzole">Sabeluzole</a></li> <li><a href="/wiki/Tabernanthine" title="Tabernanthine">Tabernanthine</a></li> <li><a href="/wiki/Tenocyclidine" title="Tenocyclidine">Tenocyclidine</a></li> <li><a href="/wiki/Tiletamine" title="Tiletamine">Tiletamine</a></li> <li><a href="/wiki/Tramadol" title="Tramadol">Tramadol</a>; Ifenprodil (NR2B) site antagonists:</li> <li><a href="/wiki/Besonprodil" title="Besonprodil">Besonprodil</a></li> <li><a href="/wiki/Buphenine" title="Buphenine">Buphenine (nylidrin)</a></li> <li><a href="/w/index.php?title=CO-101244&action=edit&redlink=1" class="new" title="CO-101244 (page does not exist)">CO-101244 (PD-174494)</a></li> <li><a href="/wiki/Eliprodil" title="Eliprodil">Eliprodil</a></li> <li><a href="/wiki/Haloperidol" title="Haloperidol">Haloperidol</a></li> <li><a href="/wiki/Isoxsuprine" title="Isoxsuprine">Isoxsuprine</a></li> <li><a href="/w/index.php?title=Radiprodil&action=edit&redlink=1" class="new" title="Radiprodil (page does not exist)">Radiprodil (RGH-896)</a></li> <li><a href="/wiki/Rislenemdaz" title="Rislenemdaz">Rislenemdaz (CERC-301, MK-0657)</a></li> <li><a href="/w/index.php?title=Ro_8-4304&action=edit&redlink=1" class="new" title="Ro 8-4304 (page does not exist)">Ro 8-4304</a></li> <li><a href="/w/index.php?title=Ro_25-6981&action=edit&redlink=1" class="new" title="Ro 25-6981 (page does not exist)">Ro 25-6981</a></li> <li><a href="/wiki/Safaprodil" class="mw-redirect" title="Safaprodil">Safaprodil</a></li> <li><a href="/wiki/Traxoprodil" title="Traxoprodil">Traxoprodil (CP-101606)</a>; NR2A-selective antagonists: <a href="/w/index.php?title=MPX-004&action=edit&redlink=1" class="new" title="MPX-004 (page does not exist)">MPX-004</a></li> <li><a href="/w/index.php?title=MPX-007&action=edit&redlink=1" class="new" title="MPX-007 (page does not exist)">MPX-007</a></li> <li><a href="/w/index.php?title=TCN-201&action=edit&redlink=1" class="new" title="TCN-201 (page does not exist)">TCN-201</a></li> <li><a href="/w/index.php?title=TCN-213&action=edit&redlink=1" class="new" title="TCN-213 (page does not exist)">TCN-213</a>; Cations: <a href="/wiki/Hydrogen" title="Hydrogen">Hydrogen</a></li> <li><a href="/wiki/Magnesium" title="Magnesium">Magnesium</a></li> <li><a href="/wiki/Zinc" title="Zinc">Zinc</a>; Alcohols/volatile anesthetics/related: <a href="/wiki/Benzene" title="Benzene">Benzene</a></li> <li><a href="/wiki/Butane" title="Butane">Butane</a></li> <li><a href="/wiki/Chloroform" title="Chloroform">Chloroform</a></li> <li><a href="/wiki/Cyclopropane" title="Cyclopropane">Cyclopropane</a></li> <li><a href="/wiki/Desflurane" title="Desflurane">Desflurane</a></li> <li><a href="/wiki/Diethyl_ether" title="Diethyl ether">Diethyl ether</a></li> <li><a href="/wiki/Enflurane" title="Enflurane">Enflurane</a></li> <li><a href="/wiki/Alcohol_(drug)" title="Alcohol (drug)">Ethanol (alcohol)</a></li> <li><a href="/wiki/Halothane" title="Halothane">Halothane</a></li> <li><a href="/wiki/Hexanol" title="Hexanol">Hexanol</a></li> <li><a href="/wiki/Isoflurane" title="Isoflurane">Isoflurane</a></li> <li><a href="/wiki/Methoxyflurane" title="Methoxyflurane">Methoxyflurane</a></li> <li><a href="/wiki/Nitrous_oxide" title="Nitrous oxide">Nitrous oxide</a></li> <li><a href="/wiki/Octanol" title="Octanol">Octanol</a></li> <li><a href="/wiki/Sevoflurane" title="Sevoflurane">Sevoflurane</a></li> <li><a href="/wiki/Toluene" title="Toluene">Toluene</a></li> <li><a href="/wiki/1,1,1-Trichloroethane" title="1,1,1-Trichloroethane">Trichloroethane</a></li> <li><a href="/wiki/2,2,2-Trichloroethanol" title="2,2,2-Trichloroethanol">Trichloroethanol</a></li> <li><a href="/wiki/Trichloroethylene" title="Trichloroethylene">Trichloroethylene</a></li> <li><a href="/wiki/Ethyl_carbamate" title="Ethyl carbamate">Urethane</a></li> <li><a href="/wiki/Xenon" title="Xenon">Xenon</a></li> <li><a href="/wiki/Xylene" title="Xylene">Xylene</a>; Unknown/unsorted antagonists: <a href="/w/index.php?title=ARR-15896&action=edit&redlink=1" class="new" title="ARR-15896 (page does not exist)">ARR-15896</a></li> <li><a href="/wiki/Bumetanide" title="Bumetanide">Bumetanide</a></li> <li><a href="/wiki/Caroverine" title="Caroverine">Caroverine</a></li> <li><a href="/wiki/Conantokin" title="Conantokin">Conantokin</a></li> <li><a href="/wiki/D-%CE%B1-Aminoadipate" class="mw-redirect" title="D-α-Aminoadipate">D-αAA</a></li> <li><a href="/wiki/Dexanabinol" title="Dexanabinol">Dexanabinol</a></li> <li><a href="/wiki/Flufenamic_acid" title="Flufenamic acid">Flufenamic acid</a></li> <li><a href="/wiki/Flupirtine" title="Flupirtine">Flupirtine</a></li> <li><a href="/w/index.php?title=FPL-12495&action=edit&redlink=1" class="new" title="FPL-12495 (page does not exist)">FPL-12495</a></li> <li><a href="/w/index.php?title=FR-115427&action=edit&redlink=1" class="new" title="FR-115427 (page does not exist)">FR-115427</a></li> <li><a href="/wiki/Furosemide" title="Furosemide">Furosemide</a></li> <li><a href="/wiki/Hodgkinsine" title="Hodgkinsine">Hodgkinsine</a></li> <li><a href="/w/index.php?title=Ipenoxazone&action=edit&redlink=1" class="new" title="Ipenoxazone (page does not exist)">Ipenoxazone (MLV-6976)</a></li> <li><a href="/w/index.php?title=MDL-27266&action=edit&redlink=1" class="new" title="MDL-27266 (page does not exist)">MDL-27266</a></li> <li><a href="/wiki/Metaphit" title="Metaphit">Metaphit</a></li> <li><a href="/wiki/Minocycline" title="Minocycline">Minocycline</a></li> <li><a href="/wiki/2-Methyl-6-(phenylethynyl)pyridine" title="2-Methyl-6-(phenylethynyl)pyridine">MPEP</a></li> <li><a href="/wiki/Niflumic_acid" title="Niflumic acid">Niflumic acid</a></li> <li><a href="/wiki/Pentamidine" title="Pentamidine">Pentamidine</a></li> <li><a href="/wiki/Pentamidine_isethionate" class="mw-redirect" title="Pentamidine isethionate">Pentamidine isethionate</a></li> <li><a href="/wiki/Piretanide" title="Piretanide">Piretanide</a></li> <li><a href="/wiki/Psychotridine" title="Psychotridine">Psychotridine</a></li> <li><a href="/wiki/Transcrocetin" class="mw-redirect" title="Transcrocetin">Transcrocetin</a> (<a href="/wiki/Saffron" title="Saffron">saffron</a>)</li></ul> <ul><li>Unsorted: Allosteric modulators: <a href="/wiki/AGN-241751" class="mw-redirect" title="AGN-241751">AGN-241751</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li>See also: <a href="/wiki/Template:Receptor_modulators" title="Template:Receptor modulators">Receptor/signaling modulators</a></li> <li><a href="/wiki/Template:Metabotropic_glutamate_receptor_modulators" title="Template:Metabotropic glutamate receptor modulators">Metabotropic glutamate receptor modulators</a></li> <li><a href="/wiki/Template:Glutamate_metabolism_and_transport_modulators" title="Template:Glutamate metabolism and transport modulators">Glutamate metabolism/transport modulators</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="Neuroethology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Neuroethology" title="Template:Neuroethology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Neuroethology" title="Template talk:Neuroethology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Neuroethology" title="Special:EditPage/Template:Neuroethology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Neuroethology" style="font-size:114%;margin:0 4em"><a href="/wiki/Neuroethology" title="Neuroethology">Neuroethology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Concepts</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/Feed_forward_(control)" title="Feed forward (control)">Feedforward</a></li> <li><a href="/wiki/Coincidence_detection_in_neurobiology" title="Coincidence detection in neurobiology">Coincidence detector</a></li> <li><a href="/wiki/Umwelt" title="Umwelt">Umwelt</a></li> <li><a href="/wiki/Instinct" title="Instinct">Instinct</a></li> <li><a href="/wiki/Feature_detection_(nervous_system)" title="Feature detection (nervous system)">Feature detection</a></li> <li><a href="/wiki/Central_pattern_generator" title="Central pattern generator">Central pattern generator (CPG)</a></li> <li><a class="mw-selflink selflink">NMDA receptor</a></li> <li><a href="/wiki/Lateral_inhibition" title="Lateral inhibition">Lateral inhibition</a></li> <li><a href="/wiki/Fixed_action_pattern" title="Fixed action pattern">Fixed action pattern</a></li> <li><a href="/wiki/Krogh%27s_Principle" class="mw-redirect" title="Krogh's Principle">Krogh's Principle</a></li> <li><a href="/wiki/Hebbian_theory" title="Hebbian theory">Hebbian theory</a></li> <li><a href="/wiki/Anti-Hebbian_learning" title="Anti-Hebbian learning">Anti-Hebbian learning</a></li> <li><a href="/wiki/Sound_localization" title="Sound localization">Sound localization</a></li> <li><a href="/wiki/Ultrasound_avoidance" title="Ultrasound avoidance">Ultrasound avoidance</a> in insects</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">People</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Theodore_Holmes_Bullock" title="Theodore Holmes Bullock">Theodore Holmes Bullock</a></li> <li><a href="/wiki/Walter_Heiligenberg" title="Walter Heiligenberg">Walter Heiligenberg</a></li> <li><a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Niko Tinbergen</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Donald_Griffin" title="Donald Griffin">Donald Griffin</a></li> <li><a href="/wiki/Donald_Kennedy" title="Donald Kennedy">Donald Kennedy</a></li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a></li> <li><a href="/wiki/Erich_von_Holst" title="Erich von Holst">Erich von Holst</a></li> <li><a href="/wiki/J%C3%B6rg-Peter_Ewert" title="Jörg-Peter Ewert">Jörg-Peter Ewert</a></li> <li><a href="/wiki/Franz_Huber" title="Franz Huber">Franz Huber</a></li> <li><a href="/wiki/Bernhard_Hassenstein" title="Bernhard Hassenstein">Bernhard Hassenstein</a></li> <li><a href="/wiki/Werner_E._Reichardt" title="Werner E. Reichardt">Werner E. Reichardt</a></li> <li><a href="/wiki/Eric_Knudsen" title="Eric Knudsen">Eric Knudsen</a></li> <li><a href="/wiki/Eric_Kandel" title="Eric Kandel">Eric Kandel</a></li> <li><a href="/wiki/Nobuo_Suga" title="Nobuo Suga">Nobuo Suga</a></li> <li><a href="/wiki/Masakazu_Konishi" title="Masakazu Konishi">Masakazu Konishi</a></li> <li><a href="/wiki/Fernando_Nottebohm" title="Fernando Nottebohm">Fernando Nottebohm</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Methods</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/Patch_clamp" title="Patch clamp">Patch clamp</a></li> <li><a href="/wiki/Slice_preparation" title="Slice preparation">Slice preparation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Systems</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_echolocation" title="Animal echolocation">Animal echolocation</a></li> <li><a href="/wiki/Waggle_dance" title="Waggle dance">Waggle dance</a></li> <li><a href="/wiki/Jamming_avoidance_response" title="Jamming avoidance response">Jamming avoidance response</a></li> <li><a href="/wiki/Vision_in_toads" title="Vision in toads">Vision in toads</a></li> <li><a href="/wiki/Frog_hearing_and_communication" title="Frog hearing and communication">Frog hearing and communication</a></li> <li><a href="/wiki/Infrared_sensing_in_snakes" title="Infrared sensing in snakes">Infrared sensing in snakes</a></li> <li><a href="/wiki/Caridoid_escape_reaction" title="Caridoid escape reaction">Caridoid escape reaction</a></li> <li><a href="/wiki/Vocal_learning" title="Vocal learning">Vocal learning</a></li> <li><a href="/wiki/Surface_wave_detection" class="mw-redirect" title="Surface wave detection">Surface wave detection</a></li> <li><a href="/wiki/Electroreception" class="mw-redirect" title="Electroreception">Electroreception</a></li> <li><a href="/wiki/Mechanoreception" class="mw-redirect" title="Mechanoreception">Mechanoreception</a> <ul><li><a href="/wiki/Lateral_line" title="Lateral line">Lateral line</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="2"><div> <ul><li><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /> <a href="/wiki/Category:Neuroethology" title="Category:Neuroethology">Category</a></li> <li><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /> <a href="https://commons.wikimedia.org/wiki/Category:Neuroethology" class="extiw" title="commons:Category:Neuroethology">Commons</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="Drug_design" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Drug_design" title="Template:Drug design"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Drug_design" title="Template talk:Drug design"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Drug_design" title="Special:EditPage/Template:Drug design"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Drug_design" style="font-size:114%;margin:0 4em"><a href="/wiki/Drug_design" title="Drug design">Drug design</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Steps in design</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Drug_discovery" title="Drug discovery">Drug discovery</a></li> <li><a href="/wiki/Hit_to_lead" title="Hit to lead">Hit to lead</a></li> <li><a href="/wiki/Drug_development" title="Drug development">Drug development</a> <ul><li><a href="/wiki/Preclinical_development" title="Preclinical development">Preclinical</a></li> <li><a href="/wiki/Clinical_trial" title="Clinical trial">Clinical</a> <ul><li><a href="/wiki/Phases_of_clinical_research" title="Phases of clinical research">Phases</a></li></ul></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Case studies of discovery and development of drug classes</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Discovery_and_development_of_5%CE%B1-reductase_inhibitors" title="Discovery and development of 5α-reductase inhibitors">5α-Reductase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_ACE_inhibitors" title="Discovery and development of ACE inhibitors">ACE inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_angiotensin_receptor_blockers" title="Discovery and development of angiotensin receptor blockers">Angiotensin receptor blockers</a></li> <li><a href="/wiki/Discovery_and_development_of_antiandrogens" title="Discovery and development of antiandrogens">Antiandrogens</a></li> <li><a href="/wiki/Discovery_and_development_of_beta-adrenergic_receptor_antagonists_(beta-blockers)" class="mw-redirect" title="Discovery and development of beta-adrenergic receptor antagonists (beta-blockers)">Beta-blockers</a></li> <li><a href="/wiki/Discovery_and_development_of_beta2_agonists" title="Discovery and development of beta2 agonists">Beta2 agonists</a></li> <li><a href="/wiki/Discovery_and_development_of_cephalosporins" title="Discovery and development of cephalosporins">Cephalosporins</a></li> <li><a href="/wiki/C-Met_inhibitors" class="mw-redirect" title="C-Met inhibitors">c-Met inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_cyclooxygenase_2_inhibitors" title="Discovery and development of cyclooxygenase 2 inhibitors">Cyclooxygenase 2 inhibitors</a></li> <li><a href="/wiki/Development_of_dipeptidyl_peptidase-4_inhibitors" class="mw-redirect" title="Development of dipeptidyl peptidase-4 inhibitors">Dipeptidyl peptidase-4 inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_direct_thrombin_inhibitors" title="Discovery and development of direct thrombin inhibitors">Direct thrombin inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_direct_Xa_inhibitors" title="Discovery and development of direct Xa inhibitors">Direct Xa inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_dual_serotonin_and_norepinephrine_reuptake_inhibitors" class="mw-redirect" title="Discovery and development of dual serotonin and norepinephrine reuptake inhibitors">Dual serotonin and norepinephrine reuptake inhibitors</a></li> <li><a href="/wiki/Development_and_discovery_of_SSRI_drugs" title="Development and discovery of SSRI drugs">Selective serotonin reuptake inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_gliflozins" title="Discovery and development of gliflozins">Gliflozins</a></li> <li><a href="/wiki/Discovery_and_development_of_HIV-protease_inhibitors" title="Discovery and development of HIV-protease inhibitors">HIV-protease inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_integrase_inhibitors" title="Discovery and development of integrase inhibitors">Integrase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_gastrointestinal_lipase_inhibitors" title="Discovery and development of gastrointestinal lipase inhibitors">Lipase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_memantine_and_related_compounds" class="mw-redirect" title="Discovery and development of memantine and related compounds">Memantine and related drugs</a></li> <li><a href="/wiki/Discovery_and_development_of_mTOR_inhibitors" class="mw-redirect" title="Discovery and development of mTOR inhibitors">mTOR inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_Neuraminidase_Inhibitors" class="mw-redirect" title="Discovery and development of Neuraminidase Inhibitors">Neuraminidase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_non-nucleoside_reverse-transcriptase_inhibitors" title="Discovery and development of non-nucleoside reverse-transcriptase inhibitors">Non-nucleoside reverse-transcriptase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_NS5A_inhibitors" title="Discovery and development of NS5A inhibitors">NS5A inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_nucleoside_and_nucleotide_reverse-transcriptase_inhibitors" title="Discovery and development of nucleoside and nucleotide reverse-transcriptase inhibitors">Nucleoside and nucleotide reverse-transcriptase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_phosphodiesterase_5_inhibitors" title="Discovery and development of phosphodiesterase 5 inhibitors">PDE5 inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_proton_pump_inhibitors" title="Discovery and development of proton pump inhibitors">Proton pump inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_statins" title="Discovery and development of statins">Statins</a></li> <li><a href="/wiki/Discovery_and_development_of_thalidomide_and_its_analogs" class="mw-redirect" title="Discovery and development of thalidomide and its analogs">Thalidomide and its analogs</a></li> <li><a href="/wiki/Discovery_and_development_of_triptans" title="Discovery and development of triptans">Triptans</a></li> <li><a href="/wiki/Discovery_and_development_of_TRPV1_antagonists" title="Discovery and development of TRPV1 antagonists">TRPV1 antagonists</a></li> <li><a href="/wiki/Discovery_and_development_of_tubulin_inhibitors" class="mw-redirect" title="Discovery and development of tubulin inhibitors">Tubulin inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_Bcr-Abl_tyrosine_kinase_inhibitors" class="mw-redirect" title="Discovery and development of Bcr-Abl tyrosine kinase inhibitors">Bcr-Abl tyrosine-kinase inhibitors</a></li> <li><a href="/wiki/Discovery_and_development_of_Cannabinoid_Receptor_1_Antagonists" class="mw-redirect" title="Discovery and development of Cannabinoid Receptor 1 Antagonists">Cannabinoid receptor antagonists</a></li> <li><a href="/wiki/Discovery_and_development_of_CCR5_receptor_antagonists" class="mw-redirect" title="Discovery and development of CCR5 receptor antagonists">CCR5 receptor antagonists</a></li> <li><a href="/wiki/Discovery_and_development_of_neurokinin_1_receptor_antagonists" class="mw-redirect" title="Discovery and development of neurokinin 1 receptor antagonists">Neurokinin 1 receptor antagonists</a></li> <li><a href="/wiki/Discovery_and_development_of_serotonin_receptor_antagonists" class="mw-redirect" title="Discovery and development of serotonin receptor antagonists">5-HT3 antagonists</a></li> <li><a href="/wiki/Discovery_and_development_of_melatonin_receptor_agonists" class="mw-redirect" title="Discovery and development of melatonin receptor agonists">Melatonin receptor agonists</a></li> <li><a href="/wiki/Discovery_and_development_of_renin_inhibitors" class="mw-redirect" title="Discovery and development of renin inhibitors">Renin inhibitors</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" 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