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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> <button aria-controls="toc-Function-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 Function subsection</span> </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Potential_function" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Potential_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Potential function</span> </div> </a> <ul id="toc-Potential_function-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Attention" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Attention"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Attention</span> </div> </a> <ul id="toc-Attention-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Empirical_evidence" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Empirical_evidence"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Empirical evidence</span> </div> </a> <ul id="toc-Empirical_evidence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Lesions_and_consciousness" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lesions_and_consciousness"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Lesions and consciousness</span> </div> </a> <ul id="toc-Lesions_and_consciousness-sublist" class="vector-toc-list"> </ul> </li> </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-Schizophrenia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Schizophrenia"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Schizophrenia</span> </div> </a> <ul id="toc-Schizophrenia-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Epilepsy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Epilepsy"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Epilepsy</span> </div> </a> <ul id="toc-Epilepsy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Consciousness" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Consciousness"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Consciousness</span> </div> </a> <ul id="toc-Consciousness-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Psilocybin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Psilocybin"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Psilocybin</span> </div> </a> <ul id="toc-Psilocybin-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Parkinsonism" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Parkinsonism"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Parkinsonism</span> </div> </a> <ul id="toc-Parkinsonism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Anxiety_and_stress" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anxiety_and_stress"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Anxiety and stress</span> </div> </a> <ul id="toc-Anxiety_and_stress-sublist" class="vector-toc-list"> </ul> </li> </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">4</span> <span>Other animals</span> </div> </a> <button aria-controls="toc-Other_animals-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 Other animals subsection</span> </button> <ul id="toc-Other_animals-sublist" class="vector-toc-list"> <li id="toc-Cats" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cats"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Cats</span> </div> </a> <ul id="toc-Cats-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Rodents" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Rodents"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Rodents</span> </div> </a> <ul id="toc-Rodents-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Monkeys" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Monkeys"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Monkeys</span> </div> </a> <ul id="toc-Monkeys-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div 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class="mw-page-title-main">Claustrum</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 19 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-19" 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">19 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B9%D8%A7%D8%A6%D9%82_(%D8%B7%D8%A8)" title="عائق (طب) – Arabic" lang="ar" hreflang="ar" data-title="عائق (طب)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Claustrum" title="Claustrum – Danish" lang="da" hreflang="da" data-title="Claustrum" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Claustrum" title="Claustrum – German" lang="de" hreflang="de" data-title="Claustrum" 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/Claustrum" title="Claustrum – Spanish" lang="es" hreflang="es" data-title="Claustrum" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Claustrum" title="Claustrum – French" lang="fr" hreflang="fr" data-title="Claustrum" 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/%EB%8B%B4%EC%9E%A5" title="담장 – Korean" lang="ko" hreflang="ko" data-title="담장" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Claustro" title="Claustro – Italian" lang="it" hreflang="it" data-title="Claustro" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A7%D7%9C%D7%90%D7%95%D7%A1%D7%98%D7%A8%D7%95%D7%9D" title="קלאוסטרום – Hebrew" lang="he" hreflang="he" data-title="קלאוסטרום" 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%9C%D0%B8_%D1%88%D0%B0%D1%80%D0%B1%D0%B0%D2%9B" title="Ми шарбақ – Kazakh" lang="kk" hreflang="kk" data-title="Ми шарбақ" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/U%C5%BEtvara" title="Užtvara – Lithuanian" lang="lt" hreflang="lt" data-title="Užtvara" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Claustrum_(hersenen)" title="Claustrum (hersenen) – Dutch" lang="nl" hreflang="nl" data-title="Claustrum (hersenen)" 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/%E5%89%8D%E9%9A%9C" title="前障 – Japanese" lang="ja" hreflang="ja" data-title="前障" 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/Przedmurze_(anatomia)" title="Przedmurze (anatomia) – Polish" lang="pl" hreflang="pl" data-title="Przedmurze (anatomia)" 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/Claustro_(sistema_nervoso)" title="Claustro (sistema nervoso) – Portuguese" lang="pt" hreflang="pt" data-title="Claustro (sistema nervoso)" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Claustru_(neuroanatomie)" title="Claustru (neuroanatomie) – Romanian" lang="ro" hreflang="ro" data-title="Claustru (neuroanatomie)" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9E%D0%B3%D1%80%D0%B0%D0%B4%D0%B0_(%D0%B0%D0%BD%D0%B0%D1%82%D0%BE%D0%BC%D0%B8%D1%8F)" title="Ограда (анатомия) – Russian" lang="ru" hreflang="ru" data-title="Ограда (анатомия)" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Claustrum" title="Claustrum – Swedish" lang="sv" hreflang="sv" data-title="Claustrum" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9E%D0%B3%D0%BE%D1%80%D0%BE%D0%B6%D0%B0_(%D0%B0%D0%BD%D0%B0%D1%82%D0%BE%D0%BC%D1%96%D1%8F)" title="Огорожа (анатомія) – Ukrainian" lang="uk" hreflang="uk" data-title="Огорожа (анатомія)" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%B1%8F%E7%8A%B6%E4%BD%93" title="屏状体 – Chinese" lang="zh" hreflang="zh" data-title="屏状体" data-language-autonym="中文" data-language-local-name="Chinese" 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<div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Structure in the brain</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Colostrum" title="Colostrum">Colostrum</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 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class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:lemonchiffon">Claustrum</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Gray_718-emphasizing-claustrum.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Gray_718-emphasizing-claustrum.png/250px-Gray_718-emphasizing-claustrum.png" decoding="async" width="250" height="268" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Gray_718-emphasizing-claustrum.png/375px-Gray_718-emphasizing-claustrum.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/50/Gray_718-emphasizing-claustrum.png/500px-Gray_718-emphasizing-claustrum.png 2x" data-file-width="550" data-file-height="590" /></a></span><div class="infobox-caption"><a href="/wiki/Coronal_section" class="mw-redirect" title="Coronal section">Coronal section</a> of human <a href="/wiki/Cerebrum" title="Cerebrum">cerebrum</a>. The claustrum is indicated by the arrow.</div></td></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Gray742-emphasizing-claustrum.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Gray742-emphasizing-claustrum.png/250px-Gray742-emphasizing-claustrum.png" decoding="async" width="250" height="286" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Gray742-emphasizing-claustrum.png/375px-Gray742-emphasizing-claustrum.png 1.5x, //upload.wikimedia.org/wikipedia/commons/d/d2/Gray742-emphasizing-claustrum.png 2x" data-file-width="500" data-file-height="572" /></a></span><div class="infobox-caption"><a href="/wiki/Transverse_section" class="mw-redirect" title="Transverse section">Transverse section</a> of human cerebrum. (Anterior is at the top.)<br />The claustrum is indicated by the arrow.</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/Brain" title="Brain">Brain</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Artery" title="Artery">Artery</a></th><td class="infobox-data"><a href="/wiki/Middle_cerebral_artery" title="Middle cerebral artery">Middle cerebral artery</a></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/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=D000079482">D000079482</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=252">252</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_1522">birnlex_1522</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/14.1.09.421%20Entity%20TA98%20EN.htm">A14.1.09.421</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=5535">5535</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%2Ffma67440">67440</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/Q429036" class="extiw" title="d:Q429036">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>The <b>claustrum</b> (Latin, meaning "to close" or "to shut") is a thin sheet of <a href="/wiki/Neuron" title="Neuron">neurons</a> and supporting <a href="/wiki/Glial_cells" class="mw-redirect" title="Glial cells">glial cells</a> in the <a href="/wiki/Brain" title="Brain">brain</a>, that connects to the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a> and subcortical regions including the <a href="/wiki/Amygdala" title="Amygdala">amygdala</a>, <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a> and <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>.<sup id="cite_ref-Crick2005_1-0" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smith1_2-0" class="reference"><a href="#cite_note-Smith1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> It is located between the <a href="/wiki/Insular_cortex" title="Insular cortex">insular cortex</a> laterally and the <a href="/wiki/Putamen" title="Putamen">putamen</a> medially, encased by the <a href="/wiki/Extreme_capsule" title="Extreme capsule">extreme</a> and <a href="/wiki/External_capsule" title="External capsule">external capsules</a> respectively.<sup id="cite_ref-Smith2019_3-0" class="reference"><a href="#cite_note-Smith2019-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Crick2005_1-1" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Chau2015_4-0" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Blood to the claustrum is supplied by the <a href="/wiki/Middle_cerebral_artery" title="Middle cerebral artery">middle cerebral artery</a>.<sup id="cite_ref-Crick2005_1-2" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> It is considered to be the most densely connected structure in the brain, and thus hypothesized to allow for the integration of various cortical inputs such as vision, sound and touch, into <a href="/wiki/Gestalt_psychology" title="Gestalt psychology">one experience</a>.<sup id="cite_ref-Chau2015_4-1" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Brown_5-0" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Other hypotheses suggest that the claustrum plays a role in <a href="/wiki/Salience_network" title="Salience network">salience processing</a>, to direct attention towards the most behaviorally relevant stimuli amongst the background noise.<sup id="cite_ref-Smith2_6-0" class="reference"><a href="#cite_note-Smith2-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The claustrum is difficult to study given the limited number of individuals with claustral lesions and the poor resolution of <a href="/wiki/Neuroimaging" title="Neuroimaging">neuroimaging</a>. </p><p>The claustrum is made up of various cell types differing in size, shape and neurochemical composition. Five cell types exist, and a majority of these cells resemble <a href="/wiki/Pyramidal_neurons" class="mw-redirect" title="Pyramidal neurons">pyramidal neurons</a> found in the cortex.<sup id="cite_ref-Braak_7-0" class="reference"><a href="#cite_note-Braak-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Nikolenko2021_8-0" class="reference"><a href="#cite_note-Nikolenko2021-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Within the claustrum, there is no <a href="/wiki/Laminar_organization" title="Laminar organization">laminar organization</a> of cell types as in the <a href="/wiki/Cerebral_cortex#Layers" title="Cerebral cortex">cortical layers</a>, and the <a href="/wiki/Soma_(biology)" title="Soma (biology)">cell bodies</a> can be a pyramidal, fusiform or circular.<sup id="cite_ref-Crick2005_1-3" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The principal cell type found in the claustrum is the <a href="/wiki/Golgi_type_I" class="mw-redirect" title="Golgi type I">Golgi type I</a> neuron, which is a large cell with <a href="/wiki/Dendrite" title="Dendrite">dendrites</a> covered in <a href="/wiki/Dendritic_spine" title="Dendritic spine">spines</a>.<sup id="cite_ref-Mathur_9-0" class="reference"><a href="#cite_note-Mathur-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Nikolenko2021_8-1" class="reference"><a href="#cite_note-Nikolenko2021-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Through interhemispheric connections, the claustrum is believed to play a role in synchronizing activity in widely separated, but functionally related, parts of the brain such as between <a href="/wiki/Frontal_eye_fields" title="Frontal eye fields">frontal eye fields</a> and the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>.<sup id="cite_ref-Crick2005_1-4" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smith2017_10-0" class="reference"><a href="#cite_note-Smith2017-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> As such, the claustrum is thought to play a role in combining different information modalities, potentially to support consciousness itself.<sup id="cite_ref-Smith2017_10-1" class="reference"><a href="#cite_note-Smith2017-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Torgerson2015_12-0" class="reference"><a href="#cite_note-Torgerson2015-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Another proposed function of the claustrum is to differentiate between relevant and irrelevant information so that the latter can be ignored.<sup id="cite_ref-Brown_5-1" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Torgerson2015_12-1" class="reference"><a href="#cite_note-Torgerson2015-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Goll2015_13-0" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Cortical components of <a href="/wiki/Consciousness" title="Consciousness">consciousness</a> include the fronto-parietal cortex, <a href="/wiki/Cingulate_cortex" title="Cingulate cortex">cingulate</a> and <a href="/wiki/Precuneus" title="Precuneus">precuneus</a>. Due to the claustrum's widespread connectivity to these areas, it is suggested that it may play a role in both <a href="/wiki/Attention" title="Attention">attention</a> and <a href="/wiki/Consciousness" title="Consciousness">consciousness</a>. The neural networks that mediate sustained attention and consciousness send inputs to the claustrum, and one case report in humans suggests that electrical stimulation near the claustrum reversibly disrupted the patient's conscious state.<sup id="cite_ref-Koubeissi-2014_14-0" class="reference"><a href="#cite_note-Koubeissi-2014-14"><span class="cite-bracket">[</span>14<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=Claustrum&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum is a small bilateral <a href="/wiki/Grey_matter" title="Grey matter">gray matter</a> structure (comprising roughly 0.25% of the cerebral cortex) located deep to the insular cortex and extreme capsule, and superficial to the external capsule and basal ganglia.<sup id="cite_ref-Crick2005_1-5" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Its name means “hidden away”, and was first identified in 1672, with more detailed descriptions coming later on during the 19th century.<sup id="cite_ref-Crick2005_1-6" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Although the regional neuroanatomical boundaries of the claustrum have been defined, there remains a lack of consensus in the literature when defining its precise margins,<sup id="cite_ref-Goll2015_13-1" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> though a meeting in 2019 of experts has posited a framework by which to refer to the structures across species.<sup id="cite_ref-Smith2019_3-1" class="reference"><a href="#cite_note-Smith2019-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Connections">Connections</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=2" title="Edit section: Connections"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An early summary of reports from the 20th century emphasized cortical inputs and outputs.<sup id="cite_ref-Edelstein2004_17-0" class="reference"><a href="#cite_note-Edelstein2004-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> However, later work has suggested the claustrum has extensive connections to cortical and subcortical regions.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> More specifically, <a href="/wiki/Electrophysiology" title="Electrophysiology">electrophysiological</a> studies show extensive connections to <a href="/wiki/List_of_thalamic_nuclei" title="List of thalamic nuclei">thalamic nuclei</a> and the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>, while isotopological reports have linked the claustrum with the prefrontal, frontal, parietal, temporal and occipital cortices.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Additional studies have also looked at the relationship of the claustrum to well-described subcortical white matter tracts. Structures such as the <a href="/wiki/Corona_radiata" title="Corona radiata">corona radiata</a>, <a href="/wiki/Occipitofrontal_fasciculus" title="Occipitofrontal fasciculus">occipitofrontal fasciculus</a> and <a href="/wiki/Uncinate_fasciculus" title="Uncinate fasciculus">uncinate fasciculus</a> project to the claustrum from frontal, pericentral, parietal and occipital regions.<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> Reciprocal connections also exist with motor, <a href="/wiki/Somatosensory_system" title="Somatosensory system">somatosensory</a>, <a href="/wiki/Primary_auditory_cortex" class="mw-redirect" title="Primary auditory cortex">auditory</a> and visual cortical regions.<sup id="cite_ref-Goll2015_13-2" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Altogether, these findings leave the claustrum as the most highly connected structure per regional volume in the brain and suggest that it may serve as a hub to coordinate activity of cerebral circuits.<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><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> Even with this extensive connectivity, most projections to and from the claustrum are ipsilateral (although there are still contralateral projections), and little evidence exists to describe its afferent or efferent connections with the brainstem and spinal cord.<sup id="cite_ref-Goll2015_13-3" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Edelstein2004_17-1" class="reference"><a href="#cite_note-Edelstein2004-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Markowitsch1984_24-0" class="reference"><a href="#cite_note-Markowitsch1984-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> In summary, the cortical and subcortical connectivity of the claustrum implies that it is most involved with processing sensory information, as well as the physical and emotional state of an animal. </p> <div class="mw-heading mw-heading3"><h3 id="Microanatomy">Microanatomy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=3" title="Edit section: Microanatomy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Inputs to the claustrum are organized by modality, which include prefrontal, visual, auditory and <a href="/wiki/Somatomotor_system" class="mw-redirect" title="Somatomotor system">somatomotor</a> processing areas. In the same way that the morphology of neurons in the <a href="/wiki/Rexed_laminae" title="Rexed laminae">Rexed laminae</a> of the <a href="/wiki/Spinal_cord" title="Spinal cord">spinal cord</a> is indicative of function, the visual, auditory and somatomotor regions within the claustrum share similar neurons with specific functional characteristics. For example, the portion of the claustrum that processes visual information (<i>primarily synthesizing afferent fibers concerned with our peripheral visual field</i>) is comprised by a majority of binocular cells that have "elongated receptive fields and no orientation selectivity".<sup id="cite_ref-Smith2010_25-0" class="reference"><a href="#cite_note-Smith2010-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>This focus on the peripheral sensory system is not an isolated occurrence, as most sensory afferents entering the claustrum bring peripheral sensory information. Moreover, the claustrum possesses a distinct <a href="/wiki/Topology" title="Topology">topological</a> organization for each sensory modality as well as the dense connectivity it shares with frontal cortices.<sup id="cite_ref-Shelton-2022_27-0" class="reference"><a href="#cite_note-Shelton-2022-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Marriott-2021_28-0" class="reference"><a href="#cite_note-Marriott-2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> For example, there is a retinotopic organization within the visual processing area of the claustrum that mirrors that of visual association cortices and V1, in a similar (yet less complicated) manner to the retinotopic conservation within the lateral geniculate nucleus.<sup id="cite_ref-Goll2015_13-4" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Within the claustrum, local connectivity is dominated by feed-forward disynaptic inhibition wherein <a href="/wiki/Parvalbumin" title="Parvalbumin">parvalbumin</a>-expressing interneurons suppress the activity of nearby projection neurons.<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> Local interneurons themselves are connected through both <a href="/wiki/Synapse" title="Synapse">chemical</a> and <a href="/wiki/Gap_junction" title="Gap junction">electrical synapses</a>, allowing for widespread and synchronous inhibition of local claustrum circuitry. In recent studies of the claustrum in <a href="/wiki/Mice" class="mw-redirect" title="Mice">mice</a><sup id="cite_ref-Shelton-2022_27-1" class="reference"><a href="#cite_note-Shelton-2022-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Bat" title="Bat">bats</a>,<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> cortically-projecting excitatory claustrum neurons were found to form synapses across the anteroposterior axis and were biased toward neurons that do not share projection targets, with the possible function of joining the activity of different afferent modules.<sup id="cite_ref-Marriott-2021_28-1" class="reference"><a href="#cite_note-Marriott-2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Combined, these two circuits suggest that the claustrum is capable of performing local transformations of diverse input information from across the brain. </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=Claustrum&action=edit&section=4" title="Edit section: Cell types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum is made up of various cell types that differ in size, shape and neurochemical composition.<sup id="cite_ref-Chau2015_4-2" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Excitatory cell types in the claustrum consist of two main classes which differentially project to <a href="/wiki/Cortex_(anatomy)" title="Cortex (anatomy)">cortical</a> and <a href="/wiki/Subcortex" class="mw-redirect" title="Subcortex">subcortical</a> brain regions.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (September 2023)">citation needed</span></a></i>]</sup> Inhibitory neurons represent only 10%-15% of the neurons within the claustrum and consist of three types, expressing <a href="/wiki/Parvalbumin" title="Parvalbumin">parvalbumin</a>, <a href="/wiki/Somatostatin_receptor_2" title="Somatostatin receptor 2">somatostatin</a> or <a href="/wiki/Vasoactive_intestinal_peptide_receptor" title="Vasoactive intestinal peptide receptor">vasoactive intestinal peptide</a>, similar to inhibitory neurons in the cortex.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Finally, many studies show that the claustrum is best distinguished structurally by its prominent plexus of parvalbumin-positive fibers formed by parvalbumin-expressing inhibitory cell types.<sup id="cite_ref-Brown_5-2" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In recent studies, the use of <a href="/wiki/Myelin_basic_protein" title="Myelin basic protein">myelin basic protein</a> (MBP) and retrogradely traveling <a href="/wiki/Cholera_toxin" title="Cholera toxin">cholera toxin</a> have additionally been used as effective methods of identifying the claustrum.<sup id="cite_ref-Marriott-2021_28-2" class="reference"><a href="#cite_note-Marriott-2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Several approaches in <a href="/wiki/Mice" class="mw-redirect" title="Mice">mice</a> have been used to assess claustrum cell types, including electrophysiological, morphological, genetic, and connectomic approaches.<sup id="cite_ref-Shelton-2022_27-2" class="reference"><a href="#cite_note-Shelton-2022-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Marriott-2021_28-3" class="reference"><a href="#cite_note-Marriott-2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Qadir-2022_33-0" class="reference"><a href="#cite_note-Qadir-2022-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><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> While no clear consensus has yet been reached regarding the exact number of excitatory cell types, recent studies have suggested that cortically- and subcortically-projecting claustrum neurons are likely distinct and vary along several metrics, such as their intrinsic electrophysiological profiles, afferent projections, and neuromodulatory profiles.<sup id="cite_ref-Shelton-2022_27-3" class="reference"><a href="#cite_note-Shelton-2022-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Qadir-2022_33-1" class="reference"><a href="#cite_note-Qadir-2022-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<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=Claustrum&action=edit&section=5" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum has been shown to have widespread activity to numerous cortical components, all of which have been associated with having components of consciousness and sustained attention. This is because of widespread connectivity to fronto-parietal areas, cingulate cortex, and thalami. Sustained attention is from the connections to the cingulate cortex, temporal cortex, and thalamus. </p><p>Crick and Koch suggest that the claustrum has a role similar to that of a conductor within an orchestra as it attempts to co-ordinate the function of all connections.<sup id="cite_ref-Crick2005_1-7" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This “conductor” analogy can also be supported through connections between the claustral, sensory, and frontal regions. The claustrum has been confirmed to be reciprocally connected to the prefrontal cortex, visual, auditory, sensory, and motor regions respectively. Connections to these modalities provide insight into the functionality of the claustrum. Here it is proposed that the claustrum functions in the gating of selective attention. Through this gating process the claustrum can selectively control input from these modalities to facilitate the process of “focusing”. It has also been suggested that it operates in the opposite context; that through divisive normalization the claustrum may implement resistance to certain input modalities to prevent “distraction”. </p> <div class="mw-heading mw-heading3"><h3 id="Potential_function">Potential function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=6" title="Edit section: Potential function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum, in order to facilitate consciousness, would need to integrate various sensory and motor modalities from various parts of the cortex. The anatomical connections of the claustrum have been observed using <a href="/wiki/Diffusion_tensor_imaging" class="mw-redirect" title="Diffusion tensor imaging">diffusion tensor imaging</a> (DTI). A <a href="/wiki/Functional_magnetic_resonance_imaging" title="Functional magnetic resonance imaging">functional magnetic resonance imaging</a> (fMRI) scan looks at oxygenated blood levels in the brain as a way of observing the activity of specific cortical areas. fMRI scans show dampened activity when anesthetized versus awake in rats, specifically claustrum connections to the <a href="/wiki/Medial_prefrontal_cortex" class="mw-redirect" title="Medial prefrontal cortex">medial prefrontal cortex</a> (mPFC) and the <a href="/wiki/Medial_dorsal_nucleus" title="Medial dorsal nucleus">mediodorsal thalamus</a> (MD thalamus). The claustrum is connected with the contralateral hemisphere's claustrum with strong, functional connections. Connections with MD thalamus, mPFC, and surrounding and distant cortical areas also exist.<sup id="cite_ref-Smith2017_10-2" class="reference"><a href="#cite_note-Smith2017-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Electrical stimulation in the dorsal claustrum of cats elicits excitatory responses within the visual cortex. The claustrum is situated anatomically at the confluence of a large number of white-matter tracts used to connect different parts of the cortex. This further suggests an integration center role for these different modalities, such as sensory and motor. Gap junctions have been shown to exist between aspiny (lacking dendrite projections) interneurons of the claustrum – suggesting a role in its ability to synchronize these modalities as input is received.<sup id="cite_ref-Crick2005_1-8" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Additional studies point to involvement in <a href="/wiki/Spatial_memory" title="Spatial memory">spatial navigation</a><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Slow-wave_sleep" title="Slow-wave sleep">slow-wave sleep</a>.<sup id="cite_ref-Norimoto2020_38-0" class="reference"><a href="#cite_note-Norimoto2020-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Narikiyo2020_39-0" class="reference"><a href="#cite_note-Narikiyo2020-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Attention">Attention</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=7" title="Edit section: Attention"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum has the differential ability to select between task-relevant information and task-irrelevant information to provide directed attention. It contains the highest density of connecting white matter tracts in the cortex. This supports the notion of networking and coordination among different regions of the brain.<sup id="cite_ref-Torgerson2015_12-2" class="reference"><a href="#cite_note-Torgerson2015-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The claustrum has regional specificity to it; information coming in from visual centers project to specific areas of grey matter neurons in the structure and the auditory cortex.<sup id="cite_ref-Crick2005_1-9" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Unexpected stimuli also activate the claustrum, effecting an immediate focusing or allocation of function. In lower mammals (e.g. rats), claustral regions receive input from somatosensory modalities, such as whiskers' motor control perspective because of its sensory and discriminatory use in these mammals.<sup id="cite_ref-Goll2015_13-5" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Functionally, it is proposed that it segregates attention between these modalities. Attention itself has been considered as top-down processing or bottom-up processing; both fit contextually with what is observed in the claustrum structurally and functionally, supporting the notion that interactions occur with high-order sensory areas involved in encoding objects and features. Input from the prefrontal cortex, for example, will define attention based upon higher-cognitive task-driven behaviour. Moreover, induction of electrical stimulation to the claustrum has been shown to cause inhibition of reading, a blank stare, and unresponsiveness. It has been reported that the claustrum has a basal frequency firing that is modulated to increase or decrease with directed attention. For example, projections to motor and oculomotor areas would assist with gaze movement to direct attention to new stimuli by increasing the firing frequency of claustral neurons.<sup id="cite_ref-Goll2015_13-6" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Salvinorin_A" title="Salvinorin A">Salvinorin A</a>, the active hallucinogenic compound found in <i><a href="/wiki/Salvia_divinorum" title="Salvia divinorum">Salvia divinorum</a></i>, is capable of inducing loss of awareness. Consumption of salvinorin A can induce synesthesia, in which different sensory modalities are interpreted by different sensory cortices. (For example: seeing sounds, tasting colours.) This supports the idea of intrathalamic segregation and conduction (attention). The claustrum has <a href="/wiki/%CE%9A-opioid_receptor" title="Κ-opioid receptor">kappa opioid receptors</a> to which Salvinorin A binds, eliciting this effect.<sup id="cite_ref-Chau2015_4-3" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Goll2015_13-7" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Empirical_evidence">Empirical evidence</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=8" title="Edit section: Empirical evidence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Tetanic_stimulation" title="Tetanic stimulation">High frequency stimulation</a> (HFS) in cat claustrum(s) has the capability to induce autonomic changes and induce “inactivation syndrome”. This syndrome is described as a decrease in awareness, indicating the relationship between the claustrum and consciousness.<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> In humans this same effect can be observed. Stimulation of the left claustrum in humans has produced "a complete arrest of volitional behavior, unresponsiveness, and amnesia without negative motor symptoms, or mere aphasia" suggesting the involvement in consciousness.<sup id="cite_ref-Koubeissi-2014_14-1" class="reference"><a href="#cite_note-Koubeissi-2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Furthermore, MRI studies have shown that increased signal intensity within the claustrum has been associated with status epilepticus – a condition in which epileptic seizures follow one another without recovery of consciousness between events.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> As well, increased signal intensity is associated with focal dyscognitive seizures, which are seizures that elicit impairment of awareness or consciousness without convulsions. The individual becomes unaware of his or her environment, and the seizure will manifest as a blank or empty stare for a window of time. </p><p>Using an operant conditioning task combined with HFS of the claustrum resulted in significant behavioural changes of rats; this included modulated motor responses, inactivity and decreased responsiveness.<sup id="cite_ref-Koubeissi-2014_14-2" class="reference"><a href="#cite_note-Koubeissi-2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Beyond this, studies have also shown that the claustrum is active during REM sleep, alongside other structures such as the dentate gyrus. These have associative roles in spatial memory, suggesting that some form of memory consolidation takes place in these areas.<sup id="cite_ref-Brown_5-3" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Lesions_and_consciousness">Lesions and consciousness</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=9" title="Edit section: Lesions and consciousness"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Functionally, the claustrum will integrate various cortical inputs through its connections into consciousness. Based upon its structure and connectivity, its function is suggested to be related to coordination of different brain functions; i.e. the conductor analogy. Consciousness functionally can be divided into two components: (i) wakefulness, which is arousal and alertness; (ii) content of consciousness, which is the processing of content. </p><p>A study of traumatic brain injuries in war veterans was undertaken to better understand the functional role of the claustrum. Damage to the claustrum was associated with the duration of one's loss of consciousness, but not its frequency. Lesion size was correlated with a longer duration of LOC events. No consequences were shown to attenuate cognitive processing.<sup id="cite_ref-Chau2015_4-4" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>In a single-case study, consciousness was shown to be disrupted when there was stimulation to the extreme capsule of the brain – which is in close proximity to the claustrum – such that upon termination of stimulation, consciousness was regained.<sup id="cite_ref-Koubeissi-2014_14-3" class="reference"><a href="#cite_note-Koubeissi-2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Another study looking at the symptomology of schizophrenia established that the severity of delusions was associated with decreased grey matter volume of the left claustrum; postulating that correlations exist between the structure and positive symptoms seen in this psychiatric disorder. Further supporting this correlation between schizophrenia and the claustrum is that there is an increase in white matter volume entering the claustrum.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Inverse correlations between grey matter volume and severity of hallucinations in the context of auditory hallucinations of schizophrenia has been supported.<sup id="cite_ref-Cascella2011_44-0" class="reference"><a href="#cite_note-Cascella2011-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> As well, to see the total loss of function of the claustrum, lesions to both claustrums on each hemisphere would need to occur.<sup id="cite_ref-Crick2005_1-10" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>However, a study in 2019, consisting of electrical stimulation of the claustrum, found no disruption of consciousness in any of the five patients that were subjected to the analysis. The tested patients reported subjective experiences in various sensory domains and exhibited reflexive movement, but none of them displayed loss of consciousness, thus questioning the claustrum's ability to disrupt consciousness when stimulated electrically.<sup id="cite_ref-Bickel2019_45-0" class="reference"><a href="#cite_note-Bickel2019-45"><span class="cite-bracket">[</span>45<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=Claustrum&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="Schizophrenia">Schizophrenia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=11" title="Edit section: Schizophrenia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Damage to the claustrum may mimic various common diseases or mental disorders; delayed development of the structure appears to be linked to <a href="/wiki/Autism" title="Autism">autism</a>. The claustrum may be involved in <a href="/wiki/Schizophrenia" title="Schizophrenia">schizophrenia</a> as findings show an increase in positive symptoms, such as delusions, when the grey matter volume of the left claustrum and right insula is decreased.<sup id="cite_ref-Cascella2011_44-1" class="reference"><a href="#cite_note-Cascella2011-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv/225px--New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="225" height="127" data-durationhint="26" data-mwtitle="New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv" data-mwprovider="wikimediacommons" resource="/wiki/File:New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv" type="video/ogg; codecs="theora, vorbis"" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="256" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/4/40/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv/New-Onset-Refractory-Status-Epilepticus-with-Claustrum-Damage-Definition-of-the-Clinical-and-Video_1.ogv.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="426" data-height="240" /></video></span><figcaption>New-onset refractory <a href="/wiki/Status_epilepticus" title="Status epilepticus">status epilepticus</a> with claustrum damage</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Epilepsy">Epilepsy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=12" title="Edit section: Epilepsy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum is also seen to play a role in epilepsy; MRIs have found increased claustral signal intensity in people who have been diagnosed with epilepsy. In certain cases, seizures tend to appear to originate from the claustrum when they are involved in early <a href="/wiki/Kainic_acid#Pharmacological_activity" title="Kainic acid">kainic acid induced seizures</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Consciousness">Consciousness</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=13" title="Edit section: Consciousness"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A single case-study showed that consciousness was disrupted when the area between the insula and claustrum was electrically stimulated; consciousness was regained when stimulation stopped.<sup id="cite_ref-Chau2015_4-5" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Koubeissi-2014_14-4" class="reference"><a href="#cite_note-Koubeissi-2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Patients that had a lesion in their left claustrum were more likely to experience a loss of consciousness compared to those that presented with lesions outside of the claustrum.<sup id="cite_ref-Chau2015_4-6" class="reference"><a href="#cite_note-Chau2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> For example, a patient that was subjected to electrode stimulation at the claustrum stopped reading, stared blankly and was unresponsive. Once the electrode was removed, the patient resumed reading and could not remember the events of being dazed.<sup id="cite_ref-Goll2015_13-8" class="reference"><a href="#cite_note-Goll2015-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>A 2019 study consisting of electrical stimulation of the claustrum found no disruption of consciousness in any of the five patients that were subjected to the analysis. The tested patients reported subjective experiences in various sensory domains and exhibited reflexive movement, but none of them displayed loss of consciousness, thus questioning the claustrum's ability to disrupt consciousness when stimulated electrically.<sup id="cite_ref-Bickel2019_45-1" class="reference"><a href="#cite_note-Bickel2019-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>A 2020 study involving artificial activation of the claustrum by optogenetic light stimulation silenced brain activity across the cortex, a phenomenon known as a "Down state," which can be seen when mice are sleeping or resting awake (quiet wakefulness).<sup id="cite_ref-Narikiyo2020_39-1" class="reference"><a href="#cite_note-Narikiyo2020-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> The authors state that 'The claustrum is a coordinator of global slow-wave activity, and it is so exciting that we are getting closer to linking specific brain connections and actions with the ultimate puzzle of consciousness.' </p><p>However, <a href="/wiki/Cognitive_science" title="Cognitive science">cognitive scientist</a> <a href="/wiki/Stevan_Harnad" title="Stevan Harnad">Stevan Harnad</a> does not consider the claustrum to be a "switch" for consciousness, but merely for wakefulness. He claims that if the claustrum were truly a switch for consciousness, artificial activation of the claustrum would not result in the patient passing out, but instead the patient would continue behaving normally, and later report that they experienced no sensations during the period of time when their claustrum was being stimulated.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Psilocybin">Psilocybin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=14" title="Edit section: Psilocybin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The claustrum expresses a high density of <a href="/wiki/5-HT2A_receptor" title="5-HT2A receptor">5HT2A</a> receptors, meaning it is significantly affected by <a href="/wiki/Serotonergic_psychedelics" class="mw-redirect" title="Serotonergic psychedelics">serotonergic psychedelics</a> like <a href="/wiki/Psilocybin" title="Psilocybin">psilocybin</a>. Psilocybin appears to affect the functional connectivity of the claustrum with the <a href="/wiki/Default_mode_network" title="Default mode network">default mode network</a> (DMN), and the fronto-parietal task control network (FPTC). Psilocybin was found to significantly decrease functional connectivity of the right claustrum with the default mode network, and increase right claustrum connectivity with the fronto-parietal task control network.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Parkinsonism">Parkinsonism</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=15" title="Edit section: Parkinsonism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A team of investigators led by neuroscientists at <a href="/wiki/Beth_Israel_Deaconess_Medical_Center" title="Beth Israel Deaconess Medical Center">Beth Israel Deaconess Medical Center</a> has identified lesions in the claustrum as the likely origin of <a href="/wiki/Parkinsonism" title="Parkinsonism">parkinsonism</a> across different conditions. The team used a novel methodology called lesion network mapping to discover the origins of parkinsonism in 29 patients whose symptoms were not the result of Parkinson's disease but rather attributed to a brain lesion – an abnormality or injury to the brain visible on brain imaging. The mapping of the 29 lesions – which were located in different regions of the brain – revealed that connectivity to the claustrum was the single most sensitive and specific marker of lesion-induced parkinsonism.<sup id="cite_ref-Joutsa2018_48-0" class="reference"><a href="#cite_note-Joutsa2018-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Anxiety_and_stress">Anxiety and stress</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=16" title="Edit section: Anxiety and stress"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In mice, suppression of claustrum appears to attenuate <a href="/wiki/Anxiety" title="Anxiety">anxiety</a>/<a href="/wiki/Psychological_stress" title="Psychological stress">stress</a> and increase <a href="/wiki/Chronic_stress" title="Chronic stress">chronic</a> stress-resistance.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </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=Claustrum&action=edit&section=17" title="Edit section: Other animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In animals, through tract tracing, findings have shown that the claustrum has extensive connections throughout the cortex with sensory and motor regions along with the hippocampus. A variety of animal models have been used such as cats, rodents and monkeys. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Anatomy_of_the_cat_(1991)_(18167720666).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg/220px-Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg" decoding="async" width="220" height="92" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg/330px-Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/39/Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg/440px-Anatomy_of_the_cat_%281991%29_%2818167720666%29.jpg 2x" data-file-width="1416" data-file-height="590" /></a><figcaption>Anatomy of a cat brain</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Cats">Cats</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=18" title="Edit section: Cats"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Technical plainlinks metadata ambox ambox-style ambox-technical" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>may be too technical for most readers to understand</b>.<span class="hide-when-compact"> Please <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Claustrum&action=edit">help improve it</a> to <a href="/wiki/Wikipedia:Make_technical_articles_understandable" title="Wikipedia:Make technical articles understandable">make it understandable to non-experts</a>, without removing the technical details.</span> <span class="date-container"><i>(<span class="date">April 2019</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>In cats, <a href="/wiki/Tetanic_stimulation" title="Tetanic stimulation">high-frequency stimulation</a> (HFS) of the claustrum can alter motor activity, induce <a href="/wiki/Autonomic_nervous_system" title="Autonomic nervous system">autonomic</a> changes, and precipitate an “inactivation syndrome” described as “decreased awareness". Recordings, primarily in cats and primates, show that claustral neurons respond to sensory stimuli and also respond during voluntary movements.<sup id="cite_ref-Brown_5-4" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Mapping from visual cortex to claustrum includes just a single map, which includes V1 and three other visual areas. Cells in the V1 are part of layer 6, and are different to cells that go to the <a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">lateral geniculate nucleus</a>; these cells use <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> as their <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a>. </p><p>The cat claustrum has 3 defined zones: (1) the anterior dorsal zone, which connects to the motor and somatosensory cortex, (2) the posterior dorsal zone that has connections to the visual cortex and (3) a third zone that is ventral to the visual one and connects to the auditory areas.<sup id="cite_ref-Crick2005_1-11" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Sensory input is segregated based on modalities and there is a high preference for peripheral sensory information. In the cat, input is received from various visual cortical areas and projects back to the area. These loops are retinotopical, meaning that regions getting visual input are responsible for the same region in the visual field as the area of the cortex that projects to the claustrum. The visual claustrum is a single map of the contralateral visual hemifield, receiving information based on motion in the visual field's periphery and has no real selectivity.<sup id="cite_ref-Olson1980_50-0" class="reference"><a href="#cite_note-Olson1980-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Sherk1981_51-0" class="reference"><a href="#cite_note-Sherk1981-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>In terms of somatosensation, cat claustrum receives dense inputs from primary somatosensory cortex (S1), but weaker inputs from secondary somatosensory cortex (S2). The inputs from S1 overlap with inputs from primary motor cortex (at least those from the forepaw representations of both).<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> Rodent claustrum does not receive input from S1 or S2, and is primarily driven by motor cortex. </p> <div class="mw-heading mw-heading3"><h3 id="Rodents">Rodents</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=19" title="Edit section: Rodents"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In rats, motor whisker areas receive input from the ipsilateral claustrum but will then project to the contralateral claustrum.<sup id="cite_ref-Brown_5-5" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The sensory barrel cortex and primary visual cortex also receive input from the ipsilateral claustrum but send very few projection back to the claustrum. Studies therefore indicate distinct patterning of connectivity of claustrum with different cortical areas. These suggest, rather than a diffuse role, they play specialized roles in cortical processing.<sup id="cite_ref-Brown_5-6" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>In mice, parvalbumin fibres are highly interconnected by chemical and electrical synapses. They are additionally also highly interconnected with claustrocortical neurons – suggesting that these inhibitory interneurons strongly modulate their activity.<sup id="cite_ref-Brown_5-7" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> These local networks suggest to synchronize activity of claustrocortical projections to therefore influence brain rhythms and co-ordinated activity of different cortical brain regions. There are additional classes of inhibitory interneurons with local connections within the claustrocortical neurons.<sup id="cite_ref-Brown_5-8" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Experiments in mice monitoring claustrocortical axonal activity to changing visual stimuli suggest the claustrum signals stimulus changes.<sup id="cite_ref-Brown_5-9" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Although claustrocortical input to visual cortical areas were engaged, the strongest responses measured were in higher-order regions of the cortex, this included the anterior cingulate cortex which is densely innervated by claustral projection.<sup id="cite_ref-Brown_5-10" class="reference"><a href="#cite_note-Brown-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Monkeys">Monkeys</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=20" title="Edit section: Monkeys"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the monkey, there are widespread connections of the claustrum with allocortical and neocortical regions. These connections project towards the frontal lobe, visual cortical regions, temporal cortex, parieto-occipital cortex and somatosensory areas amongst others.<sup id="cite_ref-Crick2005_1-12" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The subcortical areas receiving projections are the amygdala, caudate nucleus and hippocampus. It is unknown if there are cortical regions that do not receive input from the claustrum. Additionally, large or small types of aspiny neurons are reported in the monkey brain, which are classified as “local circuit neurons". </p><p>The dorsal claustrum has bi-directional connections with motor structures in the cortex.<sup id="cite_ref-Crick2005_1-13" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The relationship between the animal's movement and how neurons in the dorsocaudal claustrum behave are as follows: 70% of movement neurons are non-selective and can fire to do any push, pull or turn movements in the forelimb; the rest were more discerning and did only one of the three movements listed above.<sup id="cite_ref-Crick2005_1-14" class="reference"><a href="#cite_note-Crick2005-1"><span class="cite-bracket">[</span>1<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=Claustrum&action=edit&section=21" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Lentiform_nucleus" title="Lentiform nucleus">Lentiform nucleus</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Claustrum&action=edit&section=22" 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 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"Convergence of forepaw somatosensory and motor cortical projections in the striatum, claustrum, thalamus, and pontine nuclei of cats". <i>Brain Structure and Function</i>. <b>227</b> (1): 361–379. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00429-021-02405-6">10.1007/s00429-021-02405-6</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34665323">34665323</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:253984860">253984860</a>.</cite><span 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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/Striatum" title="Striatum">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 class="mw-selflink selflink">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 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