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class="vector-toc-list"> <li id="toc-Ventral_striatum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ventral_striatum"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Ventral striatum</span> </div> </a> <ul id="toc-Ventral_striatum-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dorsal_striatum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dorsal_striatum"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Dorsal striatum</span> </div> </a> <ul id="toc-Dorsal_striatum-sublist" class="vector-toc-list"> <li id="toc-Matrix_and_striosomes" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Matrix_and_striosomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2.1</span> <span>Matrix and striosomes</span> </div> </a> <ul id="toc-Matrix_and_striosomes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Cell_types" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cell_types"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Cell types</span> </div> </a> <ul id="toc-Cell_types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inputs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inputs"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Inputs</span> </div> </a> <ul id="toc-Inputs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Targets" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Targets"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Targets</span> </div> </a> <ul id="toc-Targets-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Blood_supply" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Blood_supply"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.6</span> <span>Blood supply</span> </div> </a> <ul id="toc-Blood_supply-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Function</span> </div> </a> <ul id="toc-Function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Clinical significance</span> </div> </a> <button aria-controls="toc-Clinical_significance-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Clinical significance subsection</span> </button> <ul id="toc-Clinical_significance-sublist" class="vector-toc-list"> <li id="toc-Parkinson's_disease_and_other_movement_disorders" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Parkinson's_disease_and_other_movement_disorders"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Parkinson's disease and other movement disorders</span> </div> </a> <ul id="toc-Parkinson's_disease_and_other_movement_disorders-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Addiction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Addiction"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Addiction</span> </div> </a> <ul id="toc-Addiction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bipolar_disorder" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bipolar_disorder"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Bipolar disorder</span> </div> </a> <ul id="toc-Bipolar_disorder-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Autism_spectrum_disorder" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Autism_spectrum_disorder"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Autism spectrum disorder</span> </div> </a> <ul id="toc-Autism_spectrum_disorder-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dysfunction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dysfunction"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Dysfunction</span> </div> </a> <ul id="toc-Dysfunction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Language_disorders" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Language_disorders"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Language disorders</span> </div> </a> <ul id="toc-Language_disorders-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_animals" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Other_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Other animals</span> </div> </a> <ul id="toc-Other_animals-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Additional_images" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Additional_images"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Additional images</span> </div> </a> <ul id="toc-Additional_images-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet 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Available in 27 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-27" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">27 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AC%D8%B3%D9%85_%D9%85%D8%AE%D8%B7%D8%B7" title="جسم مخطط – Arabic" lang="ar" hreflang="ar" data-title="جسم مخطط" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Strijatum" title="Strijatum – Bosnian" lang="bs" hreflang="bs" data-title="Strijatum" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Nucli_estriat" title="Nucli estriat – Catalan" lang="ca" hreflang="ca" data-title="Nucli estriat" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Striatum" title="Striatum – Czech" lang="cs" hreflang="cs" data-title="Striatum" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Striatum" title="Striatum – German" lang="de" hreflang="de" data-title="Striatum" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Cuerpo_estriado" title="Cuerpo estriado – Spanish" lang="es" hreflang="es" data-title="Cuerpo estriado" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Gorputz_ildaskatu" title="Gorputz ildaskatu – Basque" lang="eu" hreflang="eu" data-title="Gorputz ildaskatu" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AC%D8%B3%D9%85_%D9%85%D8%AE%D8%B7%D8%B7" title="جسم مخطط – Persian" lang="fa" hreflang="fa" data-title="جسم مخطط" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Striatum" title="Striatum – French" lang="fr" hreflang="fr" data-title="Striatum" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%84%A0%EC%A1%B0%EC%B2%B4" title="선조체 – Korean" lang="ko" hreflang="ko" data-title="선조체" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Striatum" title="Striatum – Croatian" lang="hr" hreflang="hr" data-title="Striatum" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Striatum" title="Striatum – Indonesian" lang="id" hreflang="id" data-title="Striatum" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Striato" title="Striato – Italian" lang="it" hreflang="it" data-title="Striato" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A1%D7%98%D7%A8%D7%99%D7%90%D7%98%D7%95%D7%9D" title="סטריאטום – Hebrew" lang="he" hreflang="he" data-title="סטריאטום" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%96%D0%BE%D0%BB%D0%B0%D2%9B%D1%82%D1%8B_%D0%B4%D0%B5%D0%BD%D0%B5" title="Жолақты дене – Kazakh" lang="kk" hreflang="kk" data-title="Жолақты дене" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Corpus_striatum" title="Corpus striatum – Dutch" lang="nl" hreflang="nl" data-title="Corpus striatum" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B7%9A%E6%9D%A1%E4%BD%93" title="線条体 – Japanese" lang="ja" hreflang="ja" data-title="線条体" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Pr%C4%85%C5%BCkowie" title="Prążkowie – Polish" lang="pl" hreflang="pl" data-title="Prążkowie" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Corpo_estriado" title="Corpo estriado – Portuguese" lang="pt" hreflang="pt" data-title="Corpo estriado" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%BE%D1%81%D0%B0%D1%82%D0%BE%D0%B5_%D1%82%D0%B5%D0%BB%D0%BE" title="Полосатое тело – Russian" lang="ru" hreflang="ru" data-title="Полосатое тело" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Pr%C3%BA%C5%BEkovan%C3%A9_teleso" title="Prúžkované teleso – Slovak" lang="sk" hreflang="sk" data-title="Prúžkované teleso" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Aivojuovio" title="Aivojuovio – Finnish" lang="fi" hreflang="fi" data-title="Aivojuovio" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Striatum" title="Striatum – Swedish" lang="sv" hreflang="sv" data-title="Striatum" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Striatum" title="Striatum – Turkish" lang="tr" hreflang="tr" data-title="Striatum" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D0%BC%D1%83%D0%B3%D0%B0%D1%81%D1%82%D0%B5_%D1%82%D1%96%D0%BB%D0%BE" title="Смугасте тіло – Ukrainian" lang="uk" hreflang="uk" data-title="Смугасте тіло" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%B7%9A%E6%A2%9D%E9%AB%94" title="線條體 – Cantonese" lang="yue" hreflang="yue" data-title="線條體" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%BA%B9%E7%8A%B6%E4%BD%93" title="纹状体 – Chinese" lang="zh" hreflang="zh" data-title="纹状体" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q1319792#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit 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data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">"Neostriatum" redirects here. For the avian brain region formerly called the neostriatum, see <a href="/wiki/Nidopallium" title="Nidopallium">nidopallium</a>.</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:lemonchiffon">Striatum</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg/250px-Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg" decoding="async" width="250" height="142" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg/375px-Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg/500px-Cortical_surface_with_an_overlay_of_the_basal_ganglia_and_thalamus.jpg 2x" data-file-width="1300" data-file-height="737" /></a></span><div class="infobox-caption">Striatum shown in green with other <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> and <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>. Small region in yellow is the <a href="/wiki/Amygdala" title="Amygdala">amygdala</a></div></td></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Corticostriatal_Pathway.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Corticostriatal_Pathway.jpg/250px-Corticostriatal_Pathway.jpg" decoding="async" width="250" height="237" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Corticostriatal_Pathway.jpg/375px-Corticostriatal_Pathway.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Corticostriatal_Pathway.jpg/500px-Corticostriatal_Pathway.jpg 2x" data-file-width="982" data-file-height="929" /></a></span><div class="infobox-caption"><a href="/wiki/Tractography" title="Tractography">Tractography</a> showing corticostriatal connections</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Part of</th><td class="infobox-data"><a href="/wiki/Basal_ganglia" title="Basal ganglia">Basal ganglia</a><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><br /><a href="/wiki/Reward_system" title="Reward system">Reward system</a><sup id="cite_ref-YAGER2015_2-0" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-TAYLOR2013_3-0" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Parts</th><td class="infobox-data">Ventral striatum<sup id="cite_ref-YAGER2015_2-1" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-TAYLOR2013_3-1" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-0" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><br />Dorsal striatum<sup id="cite_ref-YAGER2015_2-2" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-TAYLOR2013_3-2" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-1" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>striatum</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D003342">D003342</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/NeuroNames" title="NeuroNames">NeuroNames</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://braininfo.rprc.washington.edu/centraldirectory.aspx?ID=225">225</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/NeuroLex" title="NeuroLex">NeuroLex</a> ID</th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://uri.neuinfo.org/nif/nifstd/birnlex_1672">birnlex_1672</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Anatomica" title="Terminologia Anatomica">TA98</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/TA98%20Tree/Entity%20TA98%20EN/14.1.09.516%20Entity%20TA98%20EN.htm">A14.1.09.516</a><br /> <a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/TA98%20Tree/Entity%20TA98%20EN/14.1.09.515%20Entity%20TA98%20EN.htm">A14.1.09.515</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Anatomica" title="Terminologia Anatomica">TA2</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ta2viewer.openanatomy.org/?id=5559">5559</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma77618,">77616 77618, 77616</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_neuroanatomy" title="Anatomical terms of neuroanatomy">Anatomical terms of neuroanatomy</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q1319792" class="extiw" title="d:Q1319792">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p class="mw-empty-elt"> </p><p>The <b>striatum</b> (<abbr title="plural form">pl.</abbr>: <b>striata</b>) or <b>corpus striatum</b><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> is a cluster of interconnected <a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">nuclei</a> that make up the largest structure of the subcortical <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>.<sup id="cite_ref-Bamford_6-0" class="reference"><a href="#cite_note-Bamford-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The striatum is a critical component of the <a href="/wiki/Motor_system" title="Motor system">motor</a> and <a href="/wiki/Reward_system" title="Reward system">reward systems</a>; receives <a href="/wiki/Glutamate_(neurotransmitter)" title="Glutamate (neurotransmitter)">glutamatergic</a> and <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> inputs from different sources; and serves as the primary input to the rest of the basal ganglia. </p><p>Functionally, the striatum coordinates multiple aspects of <a href="/wiki/Cognition" title="Cognition">cognition</a>, including both <a href="/wiki/Planning#Neurology" title="Planning">motor and action planning</a>, <a href="/wiki/Decision-making" title="Decision-making">decision-making</a>, <a href="/wiki/Motivation" title="Motivation">motivation</a>, <a href="/wiki/Reinforcement" title="Reinforcement">reinforcement</a>, and <a href="/wiki/Reward_system" title="Reward system">reward</a> perception.<sup id="cite_ref-YAGER2015_2-3" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-TAYLOR2013_3-3" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-2" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The striatum is made up of the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> and the <a href="/wiki/Lentiform_nucleus" title="Lentiform nucleus">lentiform nucleus</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> However, some authors believe it is made up of <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a>, <a href="/wiki/Putamen" title="Putamen">putamen</a>, and ventral striatum.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> The lentiform nucleus is made up of the larger <a href="/wiki/Putamen" title="Putamen">putamen</a>, and the smaller <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a>.<sup id="cite_ref-Telford_10-0" class="reference"><a href="#cite_note-Telford-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Strictly speaking the globus pallidus is part of the striatum. It is common practice, however, to implicitly exclude the globus pallidus when referring to striatal structures. </p><p>In <a href="/wiki/Primate" title="Primate">primates</a>, the striatum is divided into the ventral striatum and the dorsal striatum, subdivisions that are based upon function and connections. The <a href="/wiki/Anatomical_terms_of_location#Dorsal_and_ventral" title="Anatomical terms of location">ventral</a> striatum consists of the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> and the <a href="/wiki/Olfactory_tubercle" title="Olfactory tubercle">olfactory tubercle</a>. The <a href="/wiki/Anatomical_terms_of_location#Dorsal_and_ventral" title="Anatomical terms of location">dorsal</a> striatum consists of the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> and the <a href="/wiki/Putamen" title="Putamen">putamen</a>. A <a href="/wiki/White_matter" title="White matter">white matter</a> <a href="/wiki/Nerve_tract" title="Nerve tract">nerve tract</a> (the <a href="/wiki/Internal_capsule" title="Internal capsule">internal capsule</a>) in the dorsal striatum separates the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> and the <a href="/wiki/Putamen" title="Putamen">putamen</a>.<sup id="cite_ref-FERRE2010_4-3" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Anatomically, the term <i>striatum</i> describes its striped (striated) appearance of grey-and-white matter.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Striatum_Structural_MRI.png" class="mw-file-description"><img alt="The striatum as seen on MRI. The striatum includes the caudate nucleus and the lentiform nucleus which includes the putamen and the globus pallidus" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Striatum_Structural_MRI.png/200px-Striatum_Structural_MRI.png" decoding="async" width="200" height="267" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Striatum_Structural_MRI.png/300px-Striatum_Structural_MRI.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Striatum_Structural_MRI.png/400px-Striatum_Structural_MRI.png 2x" data-file-width="577" data-file-height="769" /></a><figcaption>The striatum in red as seen on <a href="/wiki/MRI" class="mw-redirect" title="MRI">MRI</a>. The striatum includes the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> (<i>top</i>), and the <a href="/wiki/Lentiform_nucleus" title="Lentiform nucleus">lentiform nucleus</a> (the <a href="/wiki/Putamen" title="Putamen">putamen</a> (<i>right</i>) and the <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a> (<i>lower left</i>))</figcaption></figure> <p>The striatum is the largest structure of the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>. The striatum is divided into two subdivisions, a ventral striatum and a dorsal striatum, based upon function and connections. It is also divisible into a matrix and embedded striosomes. </p> <div class="mw-heading mw-heading3"><h3 id="Ventral_striatum">Ventral striatum</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=2" title="Edit section: Ventral striatum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ventral striatum is composed of the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> and the <a href="/wiki/Olfactory_tubercle" title="Olfactory tubercle">olfactory tubercle</a>.<sup id="cite_ref-FERRE2010_4-4" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid18047654_12-0" class="reference"><a href="#cite_note-pmid18047654-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The nucleus accumbens is made up of the <a href="/wiki/Nucleus_accumbens#Core" title="Nucleus accumbens">nucleus accumbens core</a> and the <a href="/wiki/Nucleus_accumbens#Shell" title="Nucleus accumbens">nucleus accumbens shell</a>, which differ by neural populations. The olfactory tubercle receives input from the <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a> but has not been shown to play a role in <a href="/wiki/Olfaction" class="mw-redirect" title="Olfaction">processing smell</a>.<sup id="cite_ref-pmid18047654_12-1" class="reference"><a href="#cite_note-pmid18047654-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> In non-primate species, the <a href="/wiki/Islands_of_Calleja" title="Islands of Calleja">islands of Calleja</a> are included.<sup id="cite_ref-:2_13-0" class="reference"><a href="#cite_note-:2-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The ventral striatum is associated with the <a href="/wiki/Limbic_system" title="Limbic system">limbic system</a> and has been implicated as a vital part of the <a href="/wiki/Neural_circuit" title="Neural circuit">circuitry</a> for decision making and reward-related behavior.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dorsal_striatum">Dorsal striatum</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=3" title="Edit section: Dorsal striatum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The dorsal striatum is composed of the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> and the <a href="/wiki/Putamen" title="Putamen">putamen</a>. Primarily it mediates cognition and involves motor and executive function. The dorsal striatum can be further subdivided into the <b>dorsomedial striatum</b>, and the <b>dorsolateral striatum</b>. Both of these areas have different roles in the acquisition of learnt behaviour and skill formation.<sup id="cite_ref-JoN2022_16-0" class="reference"><a href="#cite_note-JoN2022-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The dorsomedial region receives projections from the frontal and the parietal cortices. The dorsolateral region receives projections from the sensorimotor cortex.<sup id="cite_ref-Macpherson2019_17-0" class="reference"><a href="#cite_note-Macpherson2019-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Matrix_and_striosomes">Matrix and striosomes</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=4" title="Edit section: Matrix and striosomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Neurochemistry" title="Neurochemistry">Neurochemistry</a> studies have used <a href="/wiki/Staining" title="Staining">staining techniques</a> on the striatum that have identified two distinct striatal compartments, the matrix, and the <a href="/wiki/Striosome" title="Striosome">striosome</a> (or patch). The matrix is seen to be rich in <a href="/wiki/Acetylcholinesterase" title="Acetylcholinesterase">acetylcholinesterase</a>, while the embedded striosomes are acetylcholinesterase-poor.<sup id="cite_ref-Brimblecombe_18-0" class="reference"><a href="#cite_note-Brimblecombe-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The matrix forms the bulk of the striatum, and receives input from most areas of the cerebral cortex.<sup id="cite_ref-Squire_19-0" class="reference"><a href="#cite_note-Squire-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Clusters of neurons in the matrix, called matrisomes receive a similar input. Their output goes to both regions of the globus pallidus and to the substantia nigra pars reticulata.<sup id="cite_ref-Squire_19-1" class="reference"><a href="#cite_note-Squire-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>The striosomes receive input from the prefrontal cortex and give outputs to the substantia nigra pars compacta.<sup id="cite_ref-Squire_19-2" class="reference"><a href="#cite_note-Squire-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> There are more striosomes present in the dorsal striatum making up 10-15% of the striatal volume, than in the ventral striatum.<sup id="cite_ref-Brimblecombe_18-1" class="reference"><a href="#cite_note-Brimblecombe-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cell_types">Cell types</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=5" title="Edit section: Cell types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Dendritic_spines.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Dendritic_spines.jpg/180px-Dendritic_spines.jpg" decoding="async" width="180" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/b/b1/Dendritic_spines.jpg 1.5x" data-file-width="200" data-file-height="194" /></a><figcaption><a href="/wiki/Dendritic_spine" title="Dendritic spine">Dendritic spines</a> on <a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">medium spiny neuron</a> of striatum</figcaption></figure> <p>Types of cells in the striatum include: </p> <ul><li><a href="/wiki/Medium_spiny_neurons" class="mw-redirect" title="Medium spiny neurons">Medium spiny neurons</a> (MSNs), which are the principal neurons of the striatum.<sup id="cite_ref-YAGER2015_2-4" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> They are <a href="/wiki/GABAergic" title="GABAergic">GABAergic</a> and, thus, are classified as inhibitory neurons. Medium spiny projection neurons comprise 95% of the total neuronal population of the human striatum.<sup id="cite_ref-YAGER2015_2-5" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Medium spiny neurons have two <a href="/wiki/Phenotype" title="Phenotype">characteristic types</a>: <a href="/wiki/D1-type" class="mw-redirect" title="D1-type">D1-type</a> MSNs and <a href="/wiki/D2-type" class="mw-redirect" title="D2-type">D2-type</a> MSNs.<sup id="cite_ref-YAGER2015_2-6" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-5" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MSN_40%_mixed-type_with_DRD1_and_DRD2_20-0" class="reference"><a href="#cite_note-MSN_40%_mixed-type_with_DRD1_and_DRD2-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> A subpopulation of MSNs contain both D1-type and D2-type receptors, with approximately 40% of striatal MSNs expressing both <a href="/wiki/DRD1" class="mw-redirect" title="DRD1">DRD1</a> and <a href="/wiki/DRD2" class="mw-redirect" title="DRD2">DRD2</a> <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a>.<sup id="cite_ref-YAGER2015_2-7" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-6" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MSN_40%_mixed-type_with_DRD1_and_DRD2_20-1" class="reference"><a href="#cite_note-MSN_40%_mixed-type_with_DRD1_and_DRD2-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Acetylcholine" title="Acetylcholine">Cholinergic</a> <a href="/wiki/Interneurons" class="mw-redirect" title="Interneurons">interneurons</a> release acetylcholine, which has a variety of important effects in the striatum. In humans, other primates, and rodents, these interneurons respond to salient environmental stimuli with stereotyped responses that are temporally aligned with the responses of dopaminergic neurons of the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The large aspiny cholinergic interneurons themselves are affected by dopamine through <a href="/wiki/Dopamine_receptor_D5" title="Dopamine receptor D5">D5 dopamine receptors</a>.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Dopamine also directly controls communication between cholinergic interneurons.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li> <li>There are many types of GABAergic interneurons.<sup id="cite_ref-Ibáñez-Sandoval_O._Front_Neuroanat_2010_26-0" class="reference"><a href="#cite_note-Ibáñez-Sandoval_O._Front_Neuroanat_2010-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The best known are <a href="/wiki/Parvalbumin" title="Parvalbumin">parvalbumin</a> expressing interneurons, also known as <a href="/wiki/Action_potential" title="Action potential">fast-spiking</a> interneurons, which participate in powerful <a href="/wiki/Feed_forward_(control)" title="Feed forward (control)">feedforward</a> inhibition of principal neurons.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Also, there are GABAergic interneurons that express <a href="/wiki/Tyrosine_hydroxylase" title="Tyrosine hydroxylase">tyrosine hydroxylase</a>,<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Somatostatin" title="Somatostatin">somatostatin</a>, <a href="/wiki/Nitric_oxide_synthase" title="Nitric oxide synthase">nitric oxide synthase</a> and <a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">neuropeptide-y</a>. Recently, two types of neuropeptide-y expressing GABAergic interneurons have been described in detail,<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> one of which translates synchronous activity of cholinergic interneurons into inhibition of principal neurons.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> These <a href="/wiki/Neuron" title="Neuron">neurons</a> of the striatum are not distributed evenly.<sup id="cite_ref-Ibáñez-Sandoval_O._Front_Neuroanat_2010_26-1" class="reference"><a href="#cite_note-Ibáñez-Sandoval_O._Front_Neuroanat_2010-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup></li></ul> <p>There are two regions of <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a> in the brain – the <a href="/wiki/Subventricular_zone" title="Subventricular zone">subventricular zone</a> (SVZ) in the <a href="/wiki/Lateral_ventricle" class="mw-redirect" title="Lateral ventricle">lateral ventricles</a>, and the <a href="/wiki/Dentate_gyrus" title="Dentate gyrus">dentate gyrus</a> in the <a href="/wiki/Hippocampal_formation" title="Hippocampal formation">hippocampal formation</a>. <a href="/wiki/Neuroblast" title="Neuroblast">Neuroblasts</a> that form in the lateral ventricle adjacent to the striatum, integrate in the striatum.<sup id="cite_ref-Ernst_31-0" class="reference"><a href="#cite_note-Ernst-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> This has been noted in the human striatum following an <a href="/wiki/Stroke#Ischemic" title="Stroke">ischemic stroke</a>. Injury caused to the striatum stimulates the migration of neuroblasts from the SVZ, to the striatum, where they differentiate into adult neurons.<sup id="cite_ref-Kernie_33-0" class="reference"><a href="#cite_note-Kernie-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The normal passage of SVZ neuroblasts is to the <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a> but this traffic is diverted to the striatum after an ischemic stroke. However, few of the new developed neurons survive.<sup id="cite_ref-SVZ_34-0" class="reference"><a href="#cite_note-SVZ-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inputs">Inputs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=6" title="Edit section: Inputs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Basalganglien.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Basalganglien.png/240px-Basalganglien.png" decoding="async" width="240" height="181" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Basalganglien.png/360px-Basalganglien.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Basalganglien.png/480px-Basalganglien.png 2x" data-file-width="1594" data-file-height="1199" /></a><figcaption>Simplified diagram of frontal cortex to striatum to thalamus pathways – <a href="/wiki/Frontostriatal_circuit" title="Frontostriatal circuit">frontostriatal circuit</a></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Basal_ganglia_circuits.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/260px-Basal_ganglia_circuits.svg.png" decoding="async" width="260" height="374" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/390px-Basal_ganglia_circuits.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/520px-Basal_ganglia_circuits.svg.png 2x" data-file-width="1010" data-file-height="1453" /></a><figcaption>Overview of the main circuits of the basal ganglia. The striatum is shown in blue. Picture shows 2 coronal slices that have been superimposed to include the involved basal ganglia structures. <b>+</b> and <b>–</b> signs at the point of the arrows indicate respectively whether the pathway is excitatory or inhibitory in effect. <style data-mw-deduplicate="TemplateStyles:r1239334494">@media screen{html.skin-theme-clientpref-night .mw-parser-output div:not(.notheme)>.tmp-color,html.skin-theme-clientpref-night .mw-parser-output p>.tmp-color,html.skin-theme-clientpref-night .mw-parser-output table:not(.notheme) .tmp-color{color:inherit!important}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output div:not(.notheme)>.tmp-color,html.skin-theme-clientpref-os .mw-parser-output p>.tmp-color,html.skin-theme-clientpref-os .mw-parser-output table:not(.notheme) .tmp-color{color:inherit!important}}</style><span class="tmp-color" style="color:green">Green arrows</span> refer to excitatory <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamatergic</a> pathways, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239334494"><span class="tmp-color" style="color:red">red arrows</span> refer to inhibitory <a href="/wiki/Gamma-Aminobutyric_acid" class="mw-redirect" title="Gamma-Aminobutyric acid">GABAergic</a> pathways and <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239334494"><span class="tmp-color" style="color:turquoise">turquoise arrows</span> refer to <a href="/wiki/Dopamine" title="Dopamine">dopaminergic</a> pathways that are excitatory on the <a href="/wiki/Direct_pathway" title="Direct pathway">direct pathway</a> and inhibitory on the <a href="/wiki/Indirect_pathway" title="Indirect pathway">indirect pathway</a>.</figcaption></figure> <p>The largest connection is from the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cortex</a>, in terms of cell axons. Many parts of the <a href="/wiki/Neocortex" title="Neocortex">neocortex</a> <a href="https://en.wiktionary.org/wiki/innervate" class="extiw" title="wikt:innervate">innervate</a> the dorsal striatum. The cortical <a href="/wiki/Pyramidal_neurons" class="mw-redirect" title="Pyramidal neurons">pyramidal neurons</a> projecting to the striatum are located in layers II-VI, with the most dense projections come from layer V.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> They end mainly on the <a href="/wiki/Dendritic_spine" title="Dendritic spine">dendritic spines</a> of the spiny neurons. They are <a href="/wiki/Glutamatergic" title="Glutamatergic">glutamatergic</a>, exciting striatal neurons. </p><p>The striatum is seen as having its own internal microcircuitry.<sup id="cite_ref-Stocco_36-0" class="reference"><a href="#cite_note-Stocco-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The ventral striatum receives direct input from multiple regions in the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a> and limbic structures such as the <a href="/wiki/Amygdala" title="Amygdala">amygdala</a>, <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>, and <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, as well as the <a href="/wiki/Entorhinal_cortex" title="Entorhinal cortex">entorhinal cortex</a> and the <a href="/wiki/Inferior_temporal_gyrus" title="Inferior temporal gyrus">inferior temporal gyrus</a>.<sup id="cite_ref-:1_37-0" class="reference"><a href="#cite_note-:1-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Its primary input is to the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> system. Additionally, the <a href="/wiki/Mesolimbic_pathway" title="Mesolimbic pathway">mesolimbic pathway</a> projects from the <a href="/wiki/Ventral_tegmental_area" title="Ventral tegmental area">ventral tegmental area</a> to the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> of the ventral striatum.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p><p>Another well-known afferent is the <a href="/wiki/Nigrostriatal" class="mw-redirect" title="Nigrostriatal">nigrostriatal</a> connection arising from the neurons of the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> pars compacta. While cortical axons synapse mainly on spine heads of spiny neurons, nigral axons synapse mainly on spine shafts. In primates, the thalamostriatal afferent comes from the central median-parafascicular complex of the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> (see <a href="/wiki/Primate_basal_ganglia_system" class="mw-redirect" title="Primate basal ganglia system">primate basal ganglia system</a>). This afferent is glutamatergic. The participation of truly intralaminar neurons is much more limited. The striatum also receives afferents from other elements of the basal ganglia such as the <a href="/wiki/Subthalamic_nucleus" title="Subthalamic nucleus">subthalamic nucleus</a> (glutamatergic) or the <a href="/wiki/External_globus_pallidus" title="External globus pallidus">external globus pallidus</a> (<a href="/wiki/GABAergic" title="GABAergic">GABAergic</a>). </p> <div class="mw-heading mw-heading3"><h3 id="Targets">Targets</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=7" title="Edit section: Targets"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">Medium spiny neuron</a></div> <p>The primary outputs of the ventral striatum project to the <a href="/wiki/Ventral_pallidum" title="Ventral pallidum">ventral pallidum</a>, then the <a href="/wiki/Medial_dorsal_nucleus" title="Medial dorsal nucleus">medial dorsal nucleus</a> of the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>, which is part of the <a href="/wiki/Frontostriatal_circuit" title="Frontostriatal circuit">frontostriatal circuit</a>. Additionally, the ventral striatum projects to the <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a>, and substantia nigra pars reticulata. Some of its other outputs include projections to the <a href="/wiki/Extended_amygdala" title="Extended amygdala">extended amygdala</a>, <a href="/wiki/Lateral_hypothalamus" title="Lateral hypothalamus">lateral hypothalamus</a>, and <a href="/wiki/Pedunculopontine_nucleus" title="Pedunculopontine nucleus">pedunculopontine nucleus</a>.<sup id="cite_ref-:3_39-0" class="reference"><a href="#cite_note-:3-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>Striatal outputs from both the dorsal and ventral components are primarily composed of <a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">medium spiny neurons</a> (MSNs), a type of <a href="/wiki/Projection_neuron" class="mw-redirect" title="Projection neuron">projection neuron</a>, which have two primary <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a>: "indirect" MSNs that express <a href="/wiki/D2-like_receptor" title="D2-like receptor">D2-like receptors</a> and "direct" MSNs that express <a href="/wiki/D1-like_receptor" title="D1-like receptor">D1-like receptors</a>.<sup id="cite_ref-YAGER2015_2-8" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-7" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>The main nucleus of the basal ganglia is the striatum which projects directly to the globus pallidus via a pathway of <a href="/wiki/Striatopallidal_fibers" class="mw-redirect" title="Striatopallidal fibers">striatopallidal fibers</a>.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The striato-pallidal pathway has a whitish appearance due to the myelinated fibers. This projection comprises successively the external globus pallidus (<b>GPe</b>), the internal globus pallidus (<b>GPi</b>), the <a href="/wiki/Pars_compacta" title="Pars compacta">pars compacta</a> of the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> (<b>SNc</b>), and the <a href="/wiki/Pars_reticulata" title="Pars reticulata">pars reticulata</a> of substantia nigra (<b>SNr</b>). The neurons of this projection are inhibited by GABAergic synapses from the dorsal striatum. Among these targets, the GPe does not send axons outside the system. Others send axons to the <a href="/wiki/Superior_colliculus" title="Superior colliculus">superior colliculus</a>. Two others comprise the output to the thalamus, forming two separate channels: one through the internal segment of the globus pallidus to the ventral oralis nuclei of the thalamus and from there to the cortical <a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">supplementary motor area</a> and another through the substantia nigra to the ventral anterior nuclei of the thalamus and from there to the <a href="/wiki/Frontal_cortex" class="mw-redirect" title="Frontal cortex">frontal cortex</a> and the occulomotor cortex. </p> <div class="mw-heading mw-heading3"><h3 id="Blood_supply">Blood supply</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=8" title="Edit section: Blood supply"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Deep penetrating <a href="/wiki/Striate_arteries" class="mw-redirect" title="Striate arteries">striate arteries</a> supply blood to the striatum. These arteries include the <a href="/wiki/Recurrent_artery_of_Heubner" title="Recurrent artery of Heubner">recurrent artery of Heubner</a> arising from the <a href="/wiki/Anterior_cerebral_artery" title="Anterior cerebral artery">anterior cerebral artery</a>, and the <a href="/wiki/Lenticulostriate_arteries" class="mw-redirect" title="Lenticulostriate arteries">lenticulostriate arteries</a> arising from the <a href="/wiki/Middle_cerebral_artery" title="Middle cerebral artery">middle cerebral artery</a>.<sup id="cite_ref-Purves_41-0" class="reference"><a href="#cite_note-Purves-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=9" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ventral striatum, and the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> in particular, primarily mediates <a href="/wiki/Reward_system" title="Reward system">reward</a>, cognition, <a href="/wiki/Reinforcement" title="Reinforcement">reinforcement</a>, and <a href="/wiki/Motivational_salience" title="Motivational salience">motivational salience</a>. By contrast, the dorsal striatum primarily mediates cognition involving <a href="/wiki/Motor_function" class="mw-redirect" title="Motor function">motor function</a>, certain <a href="/wiki/Executive_functions" title="Executive functions">executive functions</a> (e.g., <a href="/wiki/Inhibitory_control" title="Inhibitory control">inhibitory control</a> and <a href="/wiki/Impulsivity" title="Impulsivity">impulsivity</a>), and <a href="/wiki/Classical_conditioning" title="Classical conditioning">stimulus-response learning</a>.<sup id="cite_ref-YAGER2015_2-9" class="reference"><a href="#cite_note-YAGER2015-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-TAYLOR2013_3-4" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FERRE2010_4-8" class="reference"><a href="#cite_note-FERRE2010-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MALENKA2009_42-0" class="reference"><a href="#cite_note-MALENKA2009-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kim_43-0" class="reference"><a href="#cite_note-Kim-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> There is a small degree of overlap, as the dorsal striatum is also a component of the <a href="/wiki/Reward_system" title="Reward system">reward system</a> that, along with the <a href="/wiki/Nucleus_accumbens_core" class="mw-redirect" title="Nucleus accumbens core">nucleus accumbens core</a>, mediates the encoding of new motor programs associated with future reward acquisition (e.g., the <a href="/wiki/Conditioned_response" class="mw-redirect" title="Conditioned response">conditioned motor response</a> to a reward cue).<sup id="cite_ref-TAYLOR2013_3-5" class="reference"><a href="#cite_note-TAYLOR2013-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MALENKA2009_42-1" class="reference"><a href="#cite_note-MALENKA2009-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>The striatum is also thought to play a role in an at least partially dissociable executive control network for language, applied to both verbal working memory and verbal attention. These models take the form of a frontal-striatal network for language processing.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> While the striatum is often not included in models of <a href="/wiki/Two-streams_hypothesis" title="Two-streams hypothesis">language processing</a>, as most models only include cortical regions, integrative models are becoming more popular in light of imaging studies, lesion studies on <a href="/wiki/Aphasia" title="Aphasia">aphasic</a> patients, and studies of language disorders concomitant with diseases known to affect the striatum like <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's</a> and <a href="/wiki/Huntington%27s_disease" title="Huntington's disease">Huntington's</a> disease.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Metabotropic" class="mw-redirect" title="Metabotropic">Metabotropic</a> <a href="/wiki/Dopamine_receptors" class="mw-redirect" title="Dopamine receptors">dopamine receptors</a> are present both on spiny neurons and on cortical axon terminals. <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">Second messenger</a> cascades triggered by activation of these dopamine receptors can modulate pre- and postsynaptic function, both in the short term and in the long term.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-DAreward_47-0" class="reference"><a href="#cite_note-DAreward-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> In humans, the striatum is activated by stimuli associated with reward, but also by <a href="/wiki/Aversive_agent" title="Aversive agent">aversive</a>, <a href="/wiki/Novelty" title="Novelty">novel</a>,<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Unexpected_hanging_paradox" title="Unexpected hanging paradox">unexpected</a>, or intense <a href="/wiki/Stimulus_(psychology)" title="Stimulus (psychology)">stimuli</a>, and cues associated with such events.<sup id="cite_ref-Volman_49-0" class="reference"><a href="#cite_note-Volman-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> <a href="/wiki/FMRI" class="mw-redirect" title="FMRI">fMRI</a> evidence suggests that the common property linking these stimuli, to which the striatum is reacting, is <a href="/wiki/Salience_(neuroscience)" title="Salience (neuroscience)">salience</a> under the conditions of presentation.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-urlDepartment_of_Physiology,_Development_and_Neuroscience:_About_the_Department_51-0" class="reference"><a href="#cite_note-urlDepartment_of_Physiology,_Development_and_Neuroscience:_About_the_Department-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> A number of other brain areas and circuits are also related to reward, such as frontal areas. Functional maps of the striatum reveal interactions with widely distributed regions of the cerebral cortex important to a diverse range of functions.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>The interplay between the striatum and the <a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">prefrontal cortex</a> is relevant for behavior, particularly adolescent development as proposed by the <a href="/wiki/Dual_systems_model" title="Dual systems model">dual systems model</a>.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_significance">Clinical significance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=10" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Parkinson's_disease_and_other_movement_disorders"><span id="Parkinson.27s_disease_and_other_movement_disorders"></span>Parkinson's disease and other movement disorders</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=11" title="Edit section: Parkinson's disease and other movement disorders"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a> results in loss of dopaminergic innervation to the dorsal striatum (and other basal ganglia) and a cascade of consequences. <a href="/wiki/Atrophy" title="Atrophy">Atrophy</a> of the striatum is also involved in <a href="/wiki/Huntington%27s_disease" title="Huntington's disease">Huntington's disease</a>, and <a href="/wiki/Movement_disorder" title="Movement disorder">movement disorders</a> such as <a href="/wiki/Chorea" title="Chorea">chorea</a>, <a href="/wiki/Choreoathetosis" title="Choreoathetosis">choreoathetosis</a>, and <a href="/wiki/Dyskinesias" class="mw-redirect" title="Dyskinesias">dyskinesias</a>.<sup id="cite_ref-lancetseminar_54-0" class="reference"><a href="#cite_note-lancetseminar-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> These have also been described as <i>circuit disorders</i> of the basal ganglia.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Addiction">Addiction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=12" title="Edit section: Addiction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Overview_of_reward_structures_in_the_human_brain.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Overview_of_reward_structures_in_the_human_brain.jpg/220px-Overview_of_reward_structures_in_the_human_brain.jpg" decoding="async" width="220" height="186" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Overview_of_reward_structures_in_the_human_brain.jpg/330px-Overview_of_reward_structures_in_the_human_brain.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Overview_of_reward_structures_in_the_human_brain.jpg/440px-Overview_of_reward_structures_in_the_human_brain.jpg 2x" data-file-width="1200" data-file-height="1017" /></a><figcaption>Overview of reward structures and associated pathways</figcaption></figure> <p><a href="/wiki/Addiction" title="Addiction">Addiction</a>, a disorder of the brain's <a href="/wiki/Reward_system" title="Reward system">reward system</a>, arises through the <a href="/wiki/Gene_expression" title="Gene expression">overexpression</a> of <a href="/wiki/FOSB#DeltaFosB" title="FOSB">DeltaFosB</a> (ΔFosB), a <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factor</a>, in the <a href="/wiki/D1-type" class="mw-redirect" title="D1-type">D1-type</a> <a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">medium spiny neurons</a> of the <a href="/wiki/Ventral_striatum" class="mw-redirect" title="Ventral striatum">ventral striatum</a>. ΔFosB is an <a href="/wiki/Inducible_gene" class="mw-redirect" title="Inducible gene">inducible gene</a> which is increasingly expressed in the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> as a result of repeatedly using an addictive drug or overexposure to other addictive stimuli.<sup id="cite_ref-Cellular_basis_56-0" class="reference"><a href="#cite_note-Cellular_basis-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Natural_and_drug_addictions_57-0" class="reference"><a href="#cite_note-Natural_and_drug_addictions-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Bipolar_disorder">Bipolar disorder</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=13" title="Edit section: Bipolar disorder"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An association has been observed between striatal expression of variants of the <a href="/wiki/PDE10A" title="PDE10A">PDE10A</a> gene and some <a href="/wiki/Bipolar_I_disorder" title="Bipolar I disorder">bipolar I disorder</a> patients. Variants of other genes, <a href="/wiki/DISC1" title="DISC1">DISC1</a> and <a href="/wiki/GNAS_complex_locus" title="GNAS complex locus">GNAS</a>, have been associated with <a href="/wiki/Bipolar_II_disorder" title="Bipolar II disorder">bipolar II disorder</a>.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Autism_spectrum_disorder">Autism spectrum disorder</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=14" title="Edit section: Autism spectrum disorder"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Autism_spectrum" class="mw-redirect" title="Autism spectrum">Autism spectrum disorder</a> (ASD) is characterized by cognitive inflexibility and poor understanding of social systems. This inflexible behavior originates in defects in the pre-frontal cortex as well as the striatal circuits.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> The defects in the striatum seem to specifically contribute to the motor, social and communication impairments seen in ASD patients. In mice which have an ASD-like phenotype induced via the overexpression of the <a href="/wiki/EIF4E" title="EIF4E">eukaryotic initiation of translation factor 4E</a>, it has been shown that these defects seem to stem from the reduced ability to store and process information in the striatum, which leads to the difficulty seen in forming new motor patterns, as well as disengaging from existing ones.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dysfunction">Dysfunction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=15" title="Edit section: Dysfunction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Dysfunction in the ventral striatum can lead to a variety of disorders, most notably <a href="/wiki/Major_depressive_disorder" title="Major depressive disorder">depression</a> and <a href="/wiki/Obsessive%E2%80%93compulsive_disorder" title="Obsessive–compulsive disorder">obsessive-compulsive disorder</a>. Because of its involvement in reward pathways, the ventral striatum has also been implicated in playing a critical role in addiction. It has been well established that the ventral striatum is strongly involved in mediating the reinforcing effects of drugs, especially stimulants, through dopaminergic stimulation.<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Language_disorders">Language disorders</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=16" title="Edit section: Language disorders"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Lesions to the striatum have been associated with deficits in speech production and comprehension. While striatal damage can impact all levels of language, damage can broadly be characterized as affecting the ability to manipulate linguistic units and rules, resulting in the promotion of default linguistic forms in conflicting situations in which selection, inhibition, and monitoring load is increased.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Two subregions of the striatum have been shown to be particularly important in language: the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a> and left <a href="/wiki/Putamen" title="Putamen">putamen</a>. Lesions localized to the caudate nucleus, as well as direct electrical stimulation, can result in <a href="/wiki/Paraphasia" title="Paraphasia">lexical paraphasias</a> and perservations (continuations of an utterance after the stimulus has ceased), which is associated with inhibited executive control, in the sense that executive control allows for the selection of the best choice among competing alternatives).<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> Stimulation of the putamen results in the inhibition of articulatory sequences and the inability to initiate motor speech commands.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=17" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the seventeenth and eighteenth centuries, the term <i>corpus striatum</i> was used to designate many distinct, deep, infracortical elements of the<sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Avoid_weasel_words" class="mw-redirect" title="Wikipedia:Avoid weasel words"><span title="The material near this tag possibly uses too vague attribution or weasel words. (April 2024)">which?</span></a></i>]</sup> hemisphere.<sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Etymologically, it is derived from (Latin) <i>striatus</i> <sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> = "grooved, striated" and the English <i>striated</i> = having parallel lines or grooves on the surface.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> In 1876 <a href="/wiki/David_Ferrier" title="David Ferrier">David Ferrier</a> contributed decades of research to the subject; concluding that the corpus striatum was vital in the "organization and generation of voluntary movement".<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> In 1941, <a href="/wiki/C%C3%A9cile_Vogt-Mugnier" title="Cécile Vogt-Mugnier">Cécile</a> and <a href="/wiki/Oskar_Vogt" title="Oskar Vogt">Oskar Vogt</a> simplified the nomenclature by proposing the term <i>striatum</i> for all elements in the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> built with striatal elements: the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a>, the <a href="/wiki/Putamen" title="Putamen">putamen</a>, and the <b>fundus striati</b>,<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> which is the ventral part linking the two preceding together ventrally to the inferior part of the <a href="/wiki/Internal_capsule" title="Internal capsule">internal capsule</a>. </p><p>The term <i><b>neostriatum</b></i> was coined by comparative anatomists comparing the subcortical structures between vertebrates, because it was thought to be a phylogenetically newer section of the corpus striatum. The term is still used by some sources, including <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a>.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Other_animals">Other animals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=18" title="Edit section: Other animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <a href="/wiki/Bird" title="Bird">birds</a> the term used was the <i>paleostriatum augmentatum</i>, and in the new avian terminology listing (as of 2002) for <i>neostriatum</i> this has been changed to the <i><a href="/wiki/Nidopallium" title="Nidopallium">nidopallium</a></i>.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p>In non-primate species, the <a href="/wiki/Islands_of_Calleja" title="Islands of Calleja">islands of Calleja</a> are included in the ventral striatum.<sup id="cite_ref-:2_13-1" class="reference"><a href="#cite_note-:2-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=19" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Cortico-basal_ganglia-thalamo-cortical_loop" title="Cortico-basal ganglia-thalamo-cortical loop">Cortico-basal ganglia-thalamo-cortical loop</a></li> <li><a href="/wiki/List_of_regions_in_the_human_brain" title="List of regions in the human brain">List of regions in the human brain</a></li> <li><a href="/wiki/Striatopallidal_fibres" title="Striatopallidal fibres">Striatopallidal fibres</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="Additional_images">Additional images</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=20" title="Edit section: Additional images"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Striatum_coronal_sections.gif" class="mw-file-description" title="Striatum highlighted in green on coronal T1 MRI images"><img alt="Striatum highlighted in green on coronal T1 MRI images" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Striatum_coronal_sections.gif/112px-Striatum_coronal_sections.gif" decoding="async" width="112" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/6/6d/Striatum_coronal_sections.gif 1.5x" data-file-width="148" data-file-height="158" /></a></span></div> <div class="gallerytext">Striatum highlighted in green on coronal T1 MRI images</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Striatum_sagittal_sections.gif" class="mw-file-description" title="Striatum highlighted in green on sagittal T1 MRI images"><img alt="Striatum highlighted in green on sagittal T1 MRI images" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/Striatum_sagittal_sections.gif/120px-Striatum_sagittal_sections.gif" decoding="async" width="120" height="102" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/Striatum_sagittal_sections.gif/180px-Striatum_sagittal_sections.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/6/62/Striatum_sagittal_sections.gif 2x" data-file-width="185" data-file-height="158" /></a></span></div> <div class="gallerytext">Striatum highlighted in green on sagittal T1 MRI images</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Striatum_transversal_sections.gif" class="mw-file-description" title="Striatum highlighted in green on transversal T1 MRI images"><img alt="Striatum highlighted in green on transversal T1 MRI images" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Striatum_transversal_sections.gif/96px-Striatum_transversal_sections.gif" decoding="async" width="96" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Striatum_transversal_sections.gif/144px-Striatum_transversal_sections.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/a/ad/Striatum_transversal_sections.gif 2x" data-file-width="148" data-file-height="185" /></a></span></div> <div class="gallerytext">Striatum highlighted in green on transversal T1 MRI images</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Striatum&action=edit&section=21" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://braininfo.rprc.washington.edu/centraldirectory.aspx?ID=224">"Basal ganglia"</a>. <i>BrainInfo</i><span class="reference-accessdate">. Retrieved <span class="nowrap">16 August</span> 2015</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=BrainInfo&rft.atitle=Basal+ganglia&rft_id=http%3A%2F%2Fbraininfo.rprc.washington.edu%2Fcentraldirectory.aspx%3FID%3D224&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStriatum" class="Z3988"></span></span> </li> <li id="cite_note-YAGER2015-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-YAGER2015_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-YAGER2015_2-9"><sup><i><b>j</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFYagerGarciaWunschFerguson2015" class="citation journal cs1">Yager LM, Garcia AF, Wunsch AM, Ferguson SM (August 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4523218">"The ins and outs of the striatum: Role in drug addiction"</a>. <i>Neuroscience</i>. <b>301</b>: 529–541. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuroscience.2015.06.033">10.1016/j.neuroscience.2015.06.033</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4523218">4523218</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26116518">26116518</a>. <q>[The striatum] receives dopaminergic inputs from the ventral tegmental area (VTA) and the substantia nigra (SNr) and glutamatergic inputs from several areas, including the cortex, hippocampus, amygdala, and thalamus (Swanson, 1982; Phillipson and Griffiths, 1985; Finch, 1996; Groenewegen et al., 1999; Britt et al., 2012). These glutamatergic inputs make contact on the heads of dendritic spines of the striatal GABAergic medium spiny projection neurons (MSNs) whereas dopaminergic inputs synapse onto the spine neck, allowing for an important and complex interaction between these two inputs in modulation of MSN activity ... It should also be noted that there is a small population of neurons in the NAc that coexpress both D1 and D2 receptors, though this is largely restricted to the NAc shell (Bertran- Gonzalez et al., 2008). ... Neurons in the NAc core and NAc shell subdivisions also differ functionally. The NAc core is involved in the processing of conditioned stimuli whereas the NAc shell is more important in the processing of unconditioned stimuli; Classically, these two striatal MSN populations are thought to have opposing effects on basal ganglia output. Activation of the dMSNs causes a net excitation of the thalamus resulting in a positive cortical feedback loop; thereby acting as a 'go' signal to initiate behavior. Activation of the iMSNs, however, causes a net inhibition of thalamic activity resulting in a negative cortical feedback loop and therefore serves as a 'brake' to inhibit behavior ... there is also mounting evidence that iMSNs play a role in motivation and addiction (Lobo and Nestler, 2011; Grueter et al., 2013). ... Together these data suggest that iMSNs normally act to restrain drug-taking behavior and recruitment of these neurons may in fact be protective against the development of compulsive drug use.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuroscience&rft.atitle=The+ins+and+outs+of+the+striatum%3A+Role+in+drug+addiction&rft.volume=301&rft.pages=529-541&rft.date=2015-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4523218%23id-name%3DPMC&rft_id=info%3Apmid%2F26116518&rft_id=info%3Adoi%2F10.1016%2Fj.neuroscience.2015.06.033&rft.aulast=Yager&rft.aufirst=LM&rft.au=Garcia%2C+AF&rft.au=Wunsch%2C+AM&rft.au=Ferguson%2C+SM&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4523218&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStriatum" class="Z3988"></span></span> </li> <li id="cite_note-TAYLOR2013-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-TAYLOR2013_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-TAYLOR2013_3-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-TAYLOR2013_3-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-TAYLOR2013_3-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-TAYLOR2013_3-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-TAYLOR2013_3-5"><sup><i><b>f</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTaylorLewisOlive2013" class="citation journal cs1">Taylor SB, Lewis CR, Olive MF (February 2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3931688">"The neurocircuitry of illicit psychostimulant addiction: acute and chronic effects in humans"</a>. <i>Subst. Abuse Rehabil</i>. <b>4</b>: 29–43. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.2147%2FSAR.S39684">10.2147/SAR.S39684</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3931688">3931688</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24648786">24648786</a>. <q>The DS (also referred to as the caudate-putamen in primates) is associated with transitions from goal-directed to habitual drug use, due in part to its role in stimulus–response learning.<sup>28,46</sup> As described above, the initial rewarding and reinforcing effects of drugs of abuse are mediated by increases in extracellular DA in the NAc shell, and after continued drug use in the NAc core.<sup>47,48</sup> After prolonged drug use, drug-associated cues produce increases in extracellular DA levels in the DS and not in the NAc.<sup>49</sup> This lends to the notion that a shift in the relative engagement from the ventral to the dorsal striatum underlies the progression from initial, voluntary drug use to habitual and compulsive drug use.<sup>28</sup> In addition to DA, recent evidence indicates that glutamatergic transmission in the DS is important for drug-induced adaptations and plasticity within the DS.<sup>50</sup></q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Subst.+Abuse+Rehabil.&rft.atitle=The+neurocircuitry+of+illicit+psychostimulant+addiction%3A+acute+and+chronic+effects+in+humans&rft.volume=4&rft.pages=29-43&rft.date=2013-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3931688%23id-name%3DPMC&rft_id=info%3Apmid%2F24648786&rft_id=info%3Adoi%2F10.2147%2FSAR.S39684&rft.aulast=Taylor&rft.aufirst=SB&rft.au=Lewis%2C+CR&rft.au=Olive%2C+MF&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3931688&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStriatum" class="Z3988"></span></span> </li> <li id="cite_note-FERRE2010-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-FERRE2010_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-FERRE2010_4-8"><sup><i><b>i</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFerréLluísJustinovaQuiroz2010" class="citation journal cs1">Ferré S, Lluís C, Justinova Z, Quiroz C, Orru M, Navarro G, Canela EI, Franco R, Goldberg SR (June 2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2931547">"Adenosine-cannabinoid receptor interactions. Implications for striatal function"</a>. <i>Br. J. Pharmacol</i>. <b>160</b> (3): 443–453. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1476-5381.2010.00723.x">10.1111/j.1476-5381.2010.00723.x</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2931547">2931547</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/20590556">20590556</a>. <q>Two classes of MSNs, which are homogeneously distributed in the striatum, can be differentiated by their output connectivity and their expression of dopamine and adenosine receptors and neuropeptides. In the dorsal striatum (mostly represented by the nucleus caudate-putamen), enkephalinergic MSNs connect the striatum with the external globus pallidus and express the peptide enkephalin and a high density of dopamine D2 and adenosine A2A receptors (they also express adenosine A1 receptors), while dynorphinergic MSNs connect the striatum with the substantia nigra (pars compacta and reticulata) and the entopeduncular nucleus (<a href="/wiki/Internal_globus_pallidus" title="Internal globus pallidus">internal globus pallidus</a>) and express the peptides dynorphin and substance P and dopamine D1 and adenosine A1 but not A2A receptors ... These two different phenotypes of MSN are also present in the ventral striatum (mostly represented by the nucleus accumbens and the olfactory tubercle). However, although they are phenotypically equal to their dorsal counterparts, they have some differences in terms of connectivity. First, not only enkephalinergic but also dynorphinergic MSNs project to the ventral counterpart of the external globus pallidus, the ventral pallidum, which, in fact, has characteristics of both the external and internal globus pallidus in its afferent and efferent connectivity. In addition to the ventral pallidum, the internal globus pallidus and the substantia nigra-VTA, the ventral striatum sends projections to the extended amygdala, the lateral hypothalamus and the pedunculopontine tegmental nucleus. ... 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Thus, acting in diverse terminal fields, dopamine confers motivational salience ("wanting") on the reward itself or associated cues (nucleus accumbens shell region), updates the value placed on different goals in light of this new experience (orbital prefrontal cortex), helps consolidate multiple forms of memory (amygdala and hippocampus), and encodes new motor programs that will facilitate obtaining this reward in the future (nucleus accumbens core region and dorsal striatum). 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.navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Rostral_basal_ganglia_and_associated_structures" title="Template:Rostral basal ganglia and associated structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Rostral_basal_ganglia_and_associated_structures" title="Template talk:Rostral basal ganglia and associated structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Rostral_basal_ganglia_and_associated_structures" title="Special:EditPage/Template:Rostral basal ganglia and associated structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Rostral_basal_ganglia_of_the_human_brain_and_associated_structures" style="font-size:114%;margin:0 4em">Rostral <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> of the <a href="/wiki/Human_brain" title="Human brain">human brain</a> and associated structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Basal_ganglia" title="Basal ganglia">Basal ganglia</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Striatum</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dorsal_striatum" class="mw-redirect" title="Dorsal striatum">Dorsal striatum</a> <ul><li><a href="/wiki/Putamen" title="Putamen">Putamen</a></li> <li><a href="/wiki/Caudate_nucleus" title="Caudate nucleus">Caudate nucleus</a></li></ul></li></ul> <ul><li><a href="/wiki/Ventral_striatum" class="mw-redirect" title="Ventral striatum">Ventral striatum</a> <ul><li><a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">Nucleus accumbens</a></li> <li><a href="/wiki/Olfactory_tubercle" title="Olfactory tubercle">Olfactory tubercle</a></li></ul></li></ul> <ul><li><a href="/wiki/Globus_pallidus" title="Globus pallidus">Globus pallidus</a> <ul><li><a href="/wiki/Lateral_globus_pallidus" class="mw-redirect" title="Lateral globus pallidus">GPe</a></li> <li><a href="/wiki/Medial_globus_pallidus" class="mw-redirect" title="Medial globus pallidus">GPi</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</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/Amygdala" title="Amygdala">Amygdala</a></li> <li><a href="/wiki/Claustrum" title="Claustrum">Claustrum</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Centrum_semiovale" title="Centrum semiovale">Centrum semiovale</a></li></ul> <ul><li><a href="/wiki/Internal_capsule" title="Internal capsule">Internal capsule</a> <ul><li><a href="/wiki/Anterior_limb_of_internal_capsule" class="mw-redirect" title="Anterior limb of internal capsule">Anterior limb</a></li> <li><a href="/wiki/Genu_of_internal_capsule" class="mw-redirect" title="Genu of internal capsule">Genu</a></li> <li><a href="/wiki/Posterior_limb_of_internal_capsule" class="mw-redirect" title="Posterior limb of internal capsule">Posterior limb</a></li> <li><a href="/wiki/Optic_radiation" title="Optic radiation">Optic radiation</a></li></ul></li></ul> <ul><li><a href="/wiki/Corona_radiata" title="Corona radiata">Corona radiata</a></li></ul> <ul><li><a href="/wiki/External_capsule" title="External capsule">External capsule</a></li> <li><a href="/wiki/Extreme_capsule" title="Extreme capsule">Extreme capsule</a></li></ul> <ul><li><i><a href="/wiki/Pallidothalamic_tracts" title="Pallidothalamic tracts">Pallidothalamic tracts</a>:</i> <a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a> <ul><li><a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a></li> <li><a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a></li></ul></li> <li><a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Rhinencephalon" title="Rhinencephalon">Rhinencephalon</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anterior_olfactory_nucleus" title="Anterior olfactory nucleus">Anterior olfactory nucleus</a></li> <li><a href="/wiki/Anterior_perforated_substance" title="Anterior perforated substance">Anterior perforated substance</a></li> <li><a href="/wiki/Olfactory_bulb" title="Olfactory bulb">Olfactory bulb</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Olfactory_tract" title="Olfactory tract">Olfactory tract</a> <ul><li><a href="/wiki/Medial_olfactory_stria" class="mw-redirect" title="Medial olfactory stria">Medial olfactory stria</a></li> <li><a href="/wiki/Lateral_olfactory_stria" class="mw-redirect" title="Lateral olfactory stria">Lateral olfactory stria</a></li></ul></li> <li><a href="/wiki/Olfactory_trigone" title="Olfactory trigone">Olfactory trigone</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other <a href="/wiki/Basal_forebrain" title="Basal forebrain">basal forebrain</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Substantia_innominata" title="Substantia innominata">Substantia innominata</a> <ul><li><a href="/wiki/Nucleus_basalis" title="Nucleus basalis">Nucleus basalis</a></li></ul></li> <li><a href="/wiki/Nucleus_of_diagonal_band" class="mw-redirect" title="Nucleus of diagonal band">Nucleus of diagonal band</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Diagonal_band_of_Broca" title="Diagonal band of Broca">Diagonal band of Broca</a></li> <li><a href="/wiki/Stria_terminalis" title="Stria terminalis">Stria terminalis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archicortex" title="Archicortex">Archicortex</a>:<br /><a href="/wiki/Hippocampal_formation" title="Hippocampal formation">Hippocampal formation</a>/<br /><a href="/wiki/Hippocampus_anatomy" title="Hippocampus anatomy">Hippocampus anatomy</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hippocampus" title="Hippocampus">Hippocampus proper</a> <ul><li><a href="/wiki/Hippocampus_proper" title="Hippocampus proper">CA1</a></li> <li><a href="/wiki/Hippocampus_proper" title="Hippocampus proper">CA2</a></li> <li><a href="/wiki/Hippocampus_proper" title="Hippocampus proper">CA3</a></li> <li><a href="/wiki/Hippocampus_proper" title="Hippocampus proper">CA4</a></li></ul></li> <li><a href="/wiki/Dentate_gyrus" title="Dentate gyrus">Dentate gyrus</a> <ul><li><a href="/wiki/Fascia_dentata" title="Fascia dentata">Fascia dentata</a></li></ul></li> <li><a href="/wiki/Subiculum" title="Subiculum">Subiculum</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alveus_of_hippocampus" class="mw-redirect" title="Alveus of hippocampus">Alveus</a></li> <li><a href="/wiki/Fimbria_of_hippocampus" class="mw-redirect" title="Fimbria of hippocampus">Fimbria</a></li> <li><a href="/wiki/Perforant_path" title="Perforant path">Perforant path</a></li> <li><a href="/wiki/Schaffer_collateral" title="Schaffer collateral">Schaffer collateral</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Brain_and_spinal_cord:_neural_tracts_and_fasciculi" 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:Neural_tracts" title="Template:Neural tracts"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Neural_tracts" title="Template talk:Neural tracts"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Neural_tracts" title="Special:EditPage/Template:Neural tracts"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Brain_and_spinal_cord:_neural_tracts_and_fasciculi" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_brain" title="Human brain">Brain</a> and <a href="/wiki/Spinal_cord" title="Spinal cord">spinal cord</a>: <a href="/wiki/Neural_pathway" title="Neural pathway">neural tracts</a> and <a href="/wiki/Nerve_fascicle" title="Nerve fascicle">fasciculi</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Sensory</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">DCML</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#First_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">1°</a>:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lamellar_corpuscle" class="mw-redirect" title="Lamellar corpuscle">Pacinian corpuscle</a>/<a href="/wiki/Tactile_corpuscle" title="Tactile corpuscle">Meissner's corpuscle</a> → <b><a href="/wiki/Posterior_column" class="mw-redirect" title="Posterior column">Posterior column</a></b> (<a href="/wiki/Gracile_fasciculus" class="mw-redirect" title="Gracile fasciculus">Gracile fasciculus</a>/<a href="/wiki/Cuneate_fasciculus" class="mw-redirect" title="Cuneate fasciculus">Cuneate fasciculus</a>) → <a href="/wiki/Gracile_nucleus" class="mw-redirect" title="Gracile nucleus">Gracile nucleus</a>/<a href="/wiki/Cuneate_nucleus" class="mw-redirect" title="Cuneate nucleus">Cuneate nucleus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#Second_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">2°</a>:</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>→ <i><a href="/wiki/Sensory_decussation" title="Sensory decussation">sensory decussation</a>/arcuate fibers</i> (<a href="/wiki/Posterior_external_arcuate_fibers" class="mw-redirect" title="Posterior external arcuate fibers">Posterior external arcuate fibers</a>, <a href="/wiki/Internal_arcuate_fibers" title="Internal arcuate fibers">Internal arcuate fibers</a>) → <a href="/wiki/Medial_lemniscus" title="Medial lemniscus">Medial lemniscus</a>/<a href="/wiki/Trigeminal_lemniscus" title="Trigeminal lemniscus">Trigeminal lemniscus</a> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_posterolateral_nucleus" title="Ventral posterolateral nucleus">VPL</a>, <a href="/wiki/Ventral_posteromedial_nucleus" title="Ventral posteromedial nucleus">VPM</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#Third_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">3°</a>:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>→ <a href="/wiki/Posterior_limb_of_internal_capsule" class="mw-redirect" title="Posterior limb of internal capsule">Posterior limb of internal capsule</a> → <a href="/wiki/Postcentral_gyrus" title="Postcentral gyrus">Postcentral gyrus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Anterolateral_system" class="mw-redirect" title="Anterolateral system">Anterolateral</a>/<br /><a href="/wiki/Pain" title="Pain">pain</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nociception#Neospinothalamic_tract" title="Nociception">Fast</a>/lateral</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1° (<a href="/wiki/Free_nerve_ending" title="Free nerve ending">Free nerve ending</a> → <a href="/wiki/A_delta_fiber" class="mw-redirect" title="A delta fiber">A delta fiber</a>) → 2° (<a href="/wiki/Anterior_white_commissure" title="Anterior white commissure">Anterior white commissure</a> → <b><a href="/wiki/Lateral_spinothalamic_tract" class="mw-redirect" title="Lateral spinothalamic tract">Lateral</a> and <a href="/wiki/Anterior_spinothalamic_tract" class="mw-redirect" title="Anterior spinothalamic tract">Anterior</a> <a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinothalamic tract</a></b> → <a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinal lemniscus</a> → <a href="/wiki/Ventral_posterolateral_nucleus" title="Ventral posterolateral nucleus">VPL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 3° (<a href="/wiki/Postcentral_gyrus" title="Postcentral gyrus">Postcentral gyrus</a>) → 4° (<a href="/wiki/Posterior_parietal_cortex" title="Posterior parietal cortex">Posterior parietal cortex</a>)</li></ul> <p>2° (<b><a href="/wiki/Spinomesencephalic_tract" class="mw-redirect" title="Spinomesencephalic tract">Spinomesencephalic tract</a></b> → <a href="/wiki/Superior_colliculus" title="Superior colliculus">Superior colliculus</a> of <a href="/wiki/Midbrain_tectum" class="mw-redirect" title="Midbrain tectum">Midbrain tectum</a>) </p> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nociception#Paleospinothalamic_tract" title="Nociception">Slow</a>/medial</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1° (<a href="/wiki/Group_C_nerve_fiber" title="Group C nerve fiber">Group C nerve fiber</a> → <b><a href="/wiki/Spinoreticular_tract" title="Spinoreticular tract">Spinoreticular tract</a></b> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a>) → 2° (<a href="/wiki/Medial_dorsal_nucleus" title="Medial dorsal nucleus">MD</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 3° (<a href="/wiki/Cingulate_cortex" title="Cingulate cortex">Cingulate cortex</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Motor</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pyramidal_tracts" title="Pyramidal tracts">Pyramidal</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i>flexion:</i> <a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a> → <a href="/wiki/Posterior_limb_of_internal_capsule" class="mw-redirect" title="Posterior limb of internal capsule">Posterior limb of internal capsule</a> → <a href="/wiki/Decussation_of_pyramids" class="mw-redirect" title="Decussation of pyramids">Decussation of pyramids</a> → <b><a href="/wiki/Pyramidal_tracts" title="Pyramidal tracts">Corticospinal tract</a> (<a href="/wiki/Lateral_corticospinal_tract" title="Lateral corticospinal tract">Lateral</a>, <a href="/wiki/Anterior_corticospinal_tract" title="Anterior corticospinal tract">Anterior</a>)</b> → <a href="/wiki/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Extrapyramidal_system" title="Extrapyramidal system">Extrapyramidal</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>flexion:</i></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a> → <a href="/wiki/Genu_of_internal_capsule" class="mw-redirect" title="Genu of internal capsule">Genu of internal capsule</a> → <b><a href="/wiki/Corticobulbar_tract" title="Corticobulbar tract">Corticobulbar tract</a></b> → <a href="/wiki/Facial_motor_nucleus" title="Facial motor nucleus">Facial motor nucleus</a> → <a href="/wiki/Facial_muscles" title="Facial muscles">Facial muscles</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>flexion:</i></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Red_nucleus" title="Red nucleus">Red nucleus</a> → <b><a href="/wiki/Rubrospinal_tract" title="Rubrospinal tract">Rubrospinal tract</a></b></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>extension:</i></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anatomy_of_the_cerebellum#Phylogenetic_and_functional_divisions" title="Anatomy of the cerebellum"> Vestibulocerebellum</a> → <a href="/wiki/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a> → <b><a href="/wiki/Vestibulospinal_tract" title="Vestibulospinal tract">Vestibulospinal tract</a></b></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>extension:</i></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anatomy_of_the_cerebellum#Phylogenetic_and_functional_divisions" title="Anatomy of the cerebellum"> Vestibulocerebellum</a> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a> → <b><a href="/wiki/Reticular_formation#Descending_reticulospinal_tracts" title="Reticular formation">Reticulospinal tract</a></b></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Midbrain_tectum" class="mw-redirect" title="Midbrain tectum">Midbrain tectum</a> → <b><a href="/wiki/Tectospinal_tract" title="Tectospinal tract">Tectospinal tract</a></b> → muscles of neck</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Basal_ganglia" title="Basal ganglia">Basal ganglia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Direct_pathway_of_movement" class="mw-redirect" title="Direct pathway of movement"><i>direct:</i></a></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">1° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a> → <a class="mw-selflink selflink">Striatum</a>) → 2° (<a href="/wiki/Medial_globus_pallidus" class="mw-redirect" title="Medial globus pallidus">GPi</a>) → 3° (<a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a>/<a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a> → <a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a> → <a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 4° (<a href="/wiki/Thalamocortical_radiations" title="Thalamocortical radiations">Thalamocortical radiations</a> → <a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor area</a>) → 5° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a>)</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Indirect_pathway_of_movement" class="mw-redirect" title="Indirect pathway of movement"><i>indirect:</i></a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">1° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a> → <a class="mw-selflink selflink">Striatum</a>) → 2° (<a href="/wiki/Lateral_globus_pallidus" class="mw-redirect" title="Lateral globus pallidus">GPe</a>) → 3° (<a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a> → <a href="/wiki/Subthalamic_nucleus" title="Subthalamic nucleus">Subthalamic nucleus</a>) → 4° (<a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a> → <a href="/wiki/Medial_globus_pallidus" class="mw-redirect" title="Medial globus pallidus">GPi</a>) → 5° (<a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a>/<a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a> → <a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a> → <a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 6° (<a href="/wiki/Thalamocortical_radiations" title="Thalamocortical radiations">Thalamocortical radiations</a> → <a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor area</a>) → 7° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a>)</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Nigrostriatal_pathway" title="Nigrostriatal pathway">nigrostriatal pathway</a>:</i></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Substantia_nigra_pars_compacta" class="mw-redirect" title="Substantia nigra pars compacta">Pars compacta</a> → <a class="mw-selflink selflink">Striatum</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cerebellum" title="Cerebellum">Cerebellar</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Afferent</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/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a> → <b><a href="/wiki/Vestibulocerebellar_tract" title="Vestibulocerebellar tract">Vestibulocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellum" title="Cerebellum">Cerebellum</a> → <a href="/wiki/Cerebellum_granule_cell" class="mw-redirect" title="Cerebellum granule cell">Granule cell</a></li></ul> <ul><li><a href="/wiki/Pontine_nuclei" title="Pontine nuclei">Pontine nuclei</a> → <b><a href="/wiki/Pontocerebellar_fibers" title="Pontocerebellar fibers">Pontocerebellar fibers</a></b> → <a href="/wiki/Middle_cerebellar_peduncle" title="Middle cerebellar peduncle">MCP</a> → <a href="/wiki/Deep_cerebellar_nuclei" title="Deep cerebellar nuclei">Deep cerebellar nuclei</a> → <a href="/wiki/Cerebellum_granule_cell" class="mw-redirect" title="Cerebellum granule cell">Granule cell</a></li></ul> <ul><li><a href="/wiki/Inferior_olivary_nucleus" title="Inferior olivary nucleus">Inferior olivary nucleus</a> → <b><a href="/wiki/Olivocerebellar_tract" title="Olivocerebellar tract">Olivocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Hemisphere</a> → <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cell</a> → <a href="/wiki/Deep_cerebellar_nuclei" title="Deep cerebellar nuclei">Deep cerebellar nuclei</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Efferent</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/Dentate_nucleus" title="Dentate nucleus">Dentate nucleus</a> in <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Lateral hemisphere/pontocerebellum</a> → <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <b><a href="/wiki/Dentatothalamic_tract" title="Dentatothalamic tract">Dentatothalamic tract</a></b> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a>) → <a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a></li></ul> <ul><li><a href="/wiki/Interposed_nucleus" title="Interposed nucleus">Interposed nucleus</a> in <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Intermediate hemisphere/spinocerebellum</a> → <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a>, or → <b><a href="/wiki/Cerebellothalamic_tract" title="Cerebellothalamic tract">Cerebellothalamic tract</a></b> → <a href="/wiki/Red_nucleus" title="Red nucleus">Red nucleus</a> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a>) → <a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a></li></ul> <ul><li><a href="/wiki/Fastigial_nucleus" title="Fastigial nucleus">Fastigial nucleus</a> in <a href="/wiki/Flocculonodular_lobe" title="Flocculonodular lobe">Flocculonodular lobe/vestibulocerebellum</a> → <b><a href="/wiki/Vestibulocerebellar_tract" title="Vestibulocerebellar tract">Vestibulocerebellar tract</a></b> → <a href="/wiki/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Bidirectional:<br /><a href="/wiki/Spinocerebellar_tract" class="mw-redirect" title="Spinocerebellar tract">Spinocerebellar</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Proprioception#Conscious_and_unconscious_proprioception" title="Proprioception">Unconscious<br /> proprioception</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lower_limb" class="mw-redirect" title="Lower limb">lower limb</a> → 1° (<a href="/wiki/Muscle_spindle" title="Muscle spindle">muscle spindles</a> → <a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">DRG</a>) → 2° (<a href="/wiki/Posterior_thoracic_nucleus" title="Posterior thoracic nucleus">Posterior thoracic nucleus</a> → <b><a href="/wiki/Dorsal_spinocerebellar_tract" class="mw-redirect" title="Dorsal spinocerebellar tract">Dorsal/posterior spinocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellar_vermis" title="Cerebellar vermis">Cerebellar vermis</a>)</li></ul> <ul><li><a href="/wiki/Upper_limb" title="Upper limb">upper limb</a> → 1° (<a href="/wiki/Muscle_spindle" title="Muscle spindle">muscle spindles</a> → <a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">DRG</a>) → 2° (<a href="/wiki/Accessory_cuneate_nucleus" title="Accessory cuneate nucleus">Accessory cuneate nucleus</a> → <b><a href="/wiki/Posterior_external_arcuate_fibers" class="mw-redirect" title="Posterior external arcuate fibers">Cuneocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Anterior_lobe_of_cerebellum" title="Anterior lobe of cerebellum">Anterior lobe of cerebellum</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Reflex_arc" title="Reflex arc">Reflex arc</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lower_limb" class="mw-redirect" title="Lower limb">lower limb</a> → 1° (<a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organ</a>) → 2° (<b><a href="/wiki/Ventral_spinocerebellar_tract" class="mw-redirect" title="Ventral spinocerebellar tract">Ventral/anterior spinocerebellar tract</a></b>→ <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <a href="/wiki/Cerebellar_vermis" title="Cerebellar vermis">Cerebellar vermis</a>)</li></ul> <ul><li><a href="/wiki/Upper_limb" title="Upper limb">upper limb</a> → 1° (<a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organ</a>) → 2° (<b><a href="/wiki/Rostral_spinocerebellar_tract" class="mw-redirect" title="Rostral spinocerebellar tract">Rostral spinocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellum" title="Cerebellum">Cerebellum</a>)</li></ul> </div></td></tr></tbody></table><div></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="Obsessive–compulsive_disorder" 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:Obsessive%E2%80%93compulsive_disorder" title="Template:Obsessive–compulsive disorder"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Obsessive%E2%80%93compulsive_disorder" title="Template talk:Obsessive–compulsive disorder"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Obsessive%E2%80%93compulsive_disorder" title="Special:EditPage/Template:Obsessive–compulsive disorder"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Obsessive–compulsive_disorder" style="font-size:114%;margin:0 4em"><a href="/wiki/Obsessive%E2%80%93compulsive_disorder" title="Obsessive–compulsive disorder">Obsessive–compulsive disorder</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">History</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/Dimensional_Obsessive-Compulsive_Scale" title="Dimensional Obsessive-Compulsive Scale">Dimensional Obsessive-Compulsive Scale</a></li> <li><a href="/wiki/Yale%E2%80%93Brown_Obsessive_Compulsive_Scale" class="mw-redirect" title="Yale–Brown Obsessive Compulsive Scale">Yale–Brown Obsessive Compulsive Scale</a></li> <li><a href="/wiki/OCD_Awareness_Week" title="OCD Awareness Week">OCD Awareness Week</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Biology_of_obsessive%E2%80%93compulsive_disorder" title="Biology of obsessive–compulsive disorder">Biology</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Neuroanatomy" scope="row" class="navbox-group" style="width:1%">Neuroanatomy</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/Basal_ganglia" title="Basal ganglia">Basal ganglia</a> (<a class="mw-selflink selflink">striatum</a>)</li> <li><a href="/wiki/Orbitofrontal_cortex" title="Orbitofrontal cortex">Orbitofrontal cortex</a></li> <li><a href="/wiki/Cingulate_cortex" title="Cingulate cortex">Cingulate cortex</a></li> <li><a href="/wiki/Brain-derived_neurotrophic_factor" title="Brain-derived neurotrophic factor">Brain-derived neurotrophic factor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Receptors</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/5-HT1D_receptor" title="5-HT1D receptor">5-HT<sub>1D</sub>β</a></li> <li><a href="/wiki/5-HT2A_receptor" title="5-HT2A receptor">5-HT<sub>2A</sub></a></li> <li><a href="/wiki/5-HT2C_receptor" title="5-HT2C receptor">5-HT<sub>2C</sub></a></li> <li><a href="/wiki/%CE%9C-opioid_receptor" title="Μ-opioid receptor">μ Opioid</a></li> <li><a href="/wiki/Histamine_H2_receptor" title="Histamine H2 receptor">H<sub>2</sub></a></li> <li><a href="/wiki/Tachykinin_receptor_1" title="Tachykinin receptor 1">NK<sub>1</sub></a></li> <li><a href="/wiki/Muscarinic_acetylcholine_receptor_M4" title="Muscarinic acetylcholine receptor M4">M<sub>4</sub></a></li> <li><a href="/wiki/NMDA_receptor" title="NMDA receptor">NMDA</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Symptoms</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/Intrusive_thoughts" class="mw-redirect" title="Intrusive thoughts">Obsessions</a> (<a href="/wiki/Magical_thinking" title="Magical thinking">associative</a></li> <li><a href="/wiki/Hypochondriasis" title="Hypochondriasis">diagnostic</a></li> <li><a href="/wiki/Self-harm" title="Self-harm">injurious</a></li> <li><a href="/wiki/Mysophobia" title="Mysophobia">pathogenic</a></li> <li><a href="/wiki/Sexual_obsessions" title="Sexual obsessions">sexual</a></li> <li><a href="/wiki/Scrupulosity" title="Scrupulosity">scrupulous</a></li> <li><a href="/wiki/Symmetry" title="Symmetry">symmetry</a>)</li> <li><a href="/wiki/Compulsive_behavior" title="Compulsive behavior">Compulsions</a> (<a href="/wiki/Impulse_(psychology)" class="mw-redirect" title="Impulse (psychology)">impulses</a></li> <li><a href="/wiki/Ritual" title="Ritual">rituals</a></li> <li><a href="/wiki/Tic" title="Tic">tics</a>)</li> <li><a href="/wiki/Thought_suppression" title="Thought suppression">Thought suppression</a> (<a href="/wiki/Experiential_avoidance" title="Experiential avoidance">Experiential avoidance</a>)</li> <li><a href="/wiki/Hoarding" title="Hoarding">Hoarding</a> (<a href="/wiki/Animal_hoarding" title="Animal hoarding">animals</a></li> <li><a href="/wiki/Bibliomania" title="Bibliomania">books</a></li> <li><a href="/wiki/Digital_hoarding" title="Digital hoarding">data</a></li> <li><a href="/wiki/Hoarding_disorder" title="Hoarding disorder">possessions</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Treatment</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="Serotonergics" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Serotonin" title="Serotonin">Serotonergics</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Selective_serotonin_reuptake_inhibitors" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Selective_serotonin_reuptake_inhibitor" title="Selective serotonin reuptake inhibitor">Selective serotonin reuptake inhibitors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Escitalopram" title="Escitalopram">Escitalopram</a></li> <li><a href="/wiki/Fluoxetine" title="Fluoxetine">Fluoxetine</a></li> <li><a href="/wiki/Fluvoxamine" title="Fluvoxamine">Fluvoxamine</a></li> <li><a href="/wiki/Paroxetine" title="Paroxetine">Paroxetine</a></li> <li><a href="/wiki/Sertraline" title="Sertraline">Sertraline</a></li> <li><a href="/wiki/Citalopram" title="Citalopram">Citalopram</a></li> <li><a href="/wiki/Nefazodone" title="Nefazodone">Nefazodone</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Monoamine_oxidase_inhibitor" title="Monoamine oxidase inhibitor">Monoamine oxidase inhibitors</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phenelzine" title="Phenelzine">Phenelzine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tricyclic_antidepressant" title="Tricyclic antidepressant">Tricyclic antidepressants</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Clomipramine" title="Clomipramine">Clomipramine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Atypical_antipsychotic" title="Atypical antipsychotic">Atypical antipsychotics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aripiprazole" title="Aripiprazole">Aripiprazole</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</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/Inositol" title="Inositol">Inositol</a></li> <li><a href="/wiki/Deep_brain_stimulation" title="Deep brain stimulation">Deep brain stimulation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Behavioral</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/Acceptance_and_commitment_therapy" title="Acceptance and commitment therapy">Acceptance and commitment therapy</a></li> <li><a href="/wiki/Cognitive_behavioral_therapy" title="Cognitive behavioral therapy">Cognitive behavioral therapy</a> (<a href="/wiki/Exposure_and_response_prevention" class="mw-redirect" title="Exposure and response prevention">Exposure and response prevention</a>)</li> <li><a href="/wiki/Inference-based_therapy" title="Inference-based therapy">Inference-based therapy</a></li> <li><a href="/wiki/Metacognitive_therapy" title="Metacognitive therapy">Metacognitive therapy</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Popular culture</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Literature</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Fictional</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Matchstick_Men" title="Matchstick Men">Matchstick Men</a></i></li> <li><i><a href="/wiki/Plyushkin" title="Plyushkin">Plyushkin</a></i></li> <li><i><a href="/wiki/Xenocide" title="Xenocide">Xenocide</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Nonfiction</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Everything_in_Its_Place" class="mw-redirect" title="Everything in Its Place">Everything in Its Place</a></i></li> <li><i><a href="/wiki/Just_Checking:_Scenes_From_the_Life_of_an_Obsessive-Compulsive" title="Just Checking: Scenes From the Life of an Obsessive-Compulsive">Just Checking</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Media</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/As_Good_as_It_Gets" title="As Good as It Gets">As Good as It Gets</a></i></li> <li><i><a href="/wiki/The_Aviator_(2004_film)" title="The Aviator (2004 film)">The Aviator</a></i></li> <li><i><a href="/wiki/Matchstick_Men" title="Matchstick Men">Matchstick Men</a></i></li> <li><i><a href="/wiki/Adrian_Monk" title="Adrian Monk">Adrian Monk</a></i></li> <li><i><a href="/wiki/Pure_(British_TV_series)" title="Pure (British TV series)">Pure</a></i></li> <li>"<a href="/wiki/$pringfield_(or,_How_I_Learned_to_Stop_Worrying_and_Love_Legalized_Gambling)" title="$pringfield (or, How I Learned to Stop Worrying and Love Legalized Gambling)">$pringfield</a>"</li> <li><i><a href="/wiki/Straight_Up_(2019_film)" title="Straight Up (2019 film)">Straight Up</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Category:Obsessive%E2%80%93compulsive_disorder" title="Category:Obsessive–compulsive disorder">Related</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Guilt_(emotion)" title="Guilt (emotion)">Guilt (emotion)</a></li> <li><a href="/wiki/Illusion_of_control" title="Illusion of control">Illusion of control</a></li> <li><a href="/wiki/Obsessive%E2%80%93compulsive_personality_disorder" title="Obsessive–compulsive personality disorder">Obsessive–compulsive personality disorder</a></li> <li><a href="/wiki/Obsessional_jealousy" title="Obsessional jealousy">Obsessional jealousy</a></li> <li><a href="/wiki/PANDAS" title="PANDAS">PANDAS</a></li> <li><a href="/wiki/Primarily_Obsessional_OCD" class="mw-redirect" title="Primarily Obsessional OCD">Primarily Obsessional OCD</a></li> <li><a href="/wiki/Questionable_cause" title="Questionable cause">Questionable cause</a></li> <li><a href="/wiki/Relationship_obsessive%E2%80%93compulsive_disorder" title="Relationship obsessive–compulsive disorder">Relationship obsessive–compulsive disorder</a></li> <li><a href="/wiki/Social_anxiety_disorder" title="Social anxiety disorder">Social anxiety disorder</a></li> <li><a href="/wiki/Superstition" title="Superstition">Superstition</a></li> <li><a href="/wiki/Tourette_syndrome" title="Tourette syndrome">Tourette syndrome</a></li> <li><a href="/wiki/University_of_Florida_Obsessive%E2%80%93Compulsive_Disorder_Program" title="University of Florida Obsessive–Compulsive Disorder Program">University of Florida Obsessive–Compulsive Disorder Program</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"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q1319792#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" 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"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q1319792#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q1319792#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Corpus striatum"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85033013">United States</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Corps strié"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb12139161w">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Corps strié"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb12139161w">BnF data</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007565363005171">Israel</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a class="external text" href="http://tools.wmflabs.org/wikidata-externalid-url/?p=1323&url_prefix=https:%2F%2Fwww.unifr.ch%2Fifaa%2FPublic%2FEntryPage%2FTA98%20Tree%2FEntity%20TA98%20EN%2F&url_suffix=%20Entity%20TA98%20EN.htm&id=A14.1.09.516">Terminologia Anatomica</a></span><ul><li><span class="uid"><a class="external text" 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