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class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Folds"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Folds</span> </div> </a> <ul id="toc-Folds-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Lobes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lobes"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Lobes</span> </div> </a> <ul id="toc-Lobes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Thickness" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Thickness"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Thickness</span> </div> </a> <ul id="toc-Thickness-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layers_of_neocortex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Layers_of_neocortex"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Layers of neocortex</span> </div> </a> <ul id="toc-Layers_of_neocortex-sublist" class="vector-toc-list"> <li id="toc-Layer_I" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_I"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.1</span> <span>Layer I</span> </div> </a> <ul id="toc-Layer_I-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_II" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_II"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.2</span> <span>Layer II</span> </div> </a> <ul id="toc-Layer_II-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_III" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_III"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.3</span> <span>Layer III</span> </div> </a> <ul id="toc-Layer_III-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_IV" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_IV"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.4</span> <span>Layer IV</span> </div> </a> <ul id="toc-Layer_IV-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_V" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_V"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.5</span> <span>Layer V</span> </div> </a> <ul id="toc-Layer_V-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_VI" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layer_VI"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.6</span> <span>Layer VI</span> </div> </a> <ul id="toc-Layer_VI-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Columns" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Columns"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Columns</span> </div> </a> <ul id="toc-Columns-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Types_of_cortex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Types_of_cortex"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.6</span> <span>Types of cortex</span> </div> </a> <ul id="toc-Types_of_cortex-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Blood_supply_and_drainage" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Blood_supply_and_drainage"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Blood supply and drainage</span> </div> </a> <ul id="toc-Blood_supply_and_drainage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Development"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Development</span> </div> </a> <button aria-controls="toc-Development-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 Development subsection</span> </button> <ul id="toc-Development-sublist" class="vector-toc-list"> <li id="toc-Neural_tube" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neural_tube"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Neural tube</span> </div> </a> <ul id="toc-Neural_tube-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cortical_neuron_development" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cortical_neuron_development"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Cortical neuron development</span> </div> </a> <ul id="toc-Cortical_neuron_development-sublist" class="vector-toc-list"> <li id="toc-Radial_glia" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Radial_glia"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Radial glia</span> </div> </a> <ul id="toc-Radial_glia-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Asymmetric_division" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Asymmetric_division"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Asymmetric division</span> </div> </a> <ul id="toc-Asymmetric_division-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Layer_organization" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Layer_organization"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Layer organization</span> </div> </a> <ul id="toc-Layer_organization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cortical_patterning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cortical_patterning"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Cortical patterning</span> </div> </a> <ul id="toc-Cortical_patterning-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</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-Connections" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Connections"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Connections</span> </div> </a> <ul id="toc-Connections-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cortical_areas" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cortical_areas"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Cortical areas</span> </div> </a> <ul id="toc-Cortical_areas-sublist" class="vector-toc-list"> <li id="toc-Sensory_areas" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Sensory_areas"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2.1</span> <span>Sensory areas</span> </div> </a> <ul id="toc-Sensory_areas-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Motor_areas" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Motor_areas"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2.2</span> <span>Motor areas</span> </div> </a> <ul id="toc-Motor_areas-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Association_areas" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Association_areas"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2.3</span> <span>Association areas</span> </div> </a> <ul id="toc-Association_areas-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</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-Genes_associated_with_cortical_disorders" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_associated_with_cortical_disorders"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Genes associated with cortical disorders</span> </div> </a> <ul id="toc-Genes_associated_with_cortical_disorders-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_animals" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Other_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Other animals</span> </div> </a> <ul id="toc-Other_animals-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Additional_images" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Additional_images"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Additional images</span> </div> </a> <ul id="toc-Additional_images-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</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"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Cerebral cortex</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 52 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-52" 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">52 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%82%D8%B4%D8%B1%D8%A9_%D9%85%D8%AE%D9%8A%D8%A9" title="قشرة مخية – Arabic" lang="ar" hreflang="ar" data-title="قشرة مخية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Corteya_cerebral" title="Corteya cerebral – Asturian" lang="ast" hreflang="ast" data-title="Corteya cerebral" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%D8%A8%D8%A6%DB%8C%DB%8C%D9%86_%DA%A9%D9%88%D8%B1%D8%AA%DA%A9%D8%B3%DB%8C" title="بئیین کورتکسی – South Azerbaijani" lang="azb" hreflang="azb" data-title="بئیین کورتکسی" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target"><span>تۆرکجه</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9A%D0%B0%D1%80%D0%B0_%D0%B2%D1%8F%D0%BB%D1%96%D0%BA%D1%96%D1%85_%D0%BF%D0%B0%D1%9E%D1%88%D0%B0%D1%80%E2%80%99%D1%8F%D1%9E" title="Кара вялікіх паўшар’яў – Belarusian" lang="be" hreflang="be" data-title="Кара вялікіх паўшар’яў" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9C%D0%BE%D0%B7%D1%8A%D1%87%D0%BD%D0%B0_%D0%BA%D0%BE%D1%80%D0%B0" title="Мозъчна кора – Bulgarian" lang="bg" hreflang="bg" data-title="Мозъчна кора" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Mo%C5%BEdana_kora" title="Moždana kora – Bosnian" lang="bs" hreflang="bs" data-title="Moždana kora" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Escor%C3%A7a_cerebral" title="Escorça cerebral – Catalan" lang="ca" hreflang="ca" data-title="Escorça cerebral" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Mozkov%C3%A1_k%C5%AFra" title="Mozková kůra – Czech" lang="cs" hreflang="cs" data-title="Mozková kůra" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Hjernebark" title="Hjernebark – Danish" lang="da" hreflang="da" data-title="Hjernebark" 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/Gro%C3%9Fhirnrinde" title="Großhirnrinde – German" lang="de" hreflang="de" data-title="Großhirnrinde" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Suurajukoor" title="Suurajukoor – Estonian" lang="et" hreflang="et" data-title="Suurajukoor" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%95%CE%B3%CE%BA%CE%B5%CF%86%CE%B1%CE%BB%CE%B9%CE%BA%CF%8C%CF%82_%CF%86%CE%BB%CE%BF%CE%B9%CF%8C%CF%82" title="Εγκεφαλικός φλοιός – Greek" lang="el" hreflang="el" data-title="Εγκεφαλικός φλοιός" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Corteza_cerebral" title="Corteza cerebral – Spanish" lang="es" hreflang="es" data-title="Corteza cerebral" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Kortekso" title="Kortekso – Esperanto" lang="eo" hreflang="eo" data-title="Kortekso" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Kortex_zerebral" title="Kortex zerebral – Basque" lang="eu" hreflang="eu" data-title="Kortex zerebral" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%82%D8%B4%D8%B1_%D9%85%D8%BA%D8%B2" title="قشر مغز – Persian" lang="fa" hreflang="fa" data-title="قشر مغز" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Cortex_c%C3%A9r%C3%A9bral" title="Cortex cérébral – French" lang="fr" hreflang="fr" data-title="Cortex cérébral" 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%8C%80%EB%87%8C_%ED%94%BC%EC%A7%88" title="대뇌 피질 – Korean" lang="ko" hreflang="ko" data-title="대뇌 피질" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%84%D5%A5%D5%AE_%D5%B8%D6%82%D5%B2%D5%A5%D5%B2%D5%AB_%D5%AF%D5%A5%D5%B2%D6%87" title="Մեծ ուղեղի կեղև – Armenian" lang="hy" hreflang="hy" data-title="Մեծ ուղեղի կեղև" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Korteks_otak_besar" title="Korteks otak besar – Indonesian" lang="id" hreflang="id" data-title="Korteks otak besar" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Corteccia_cerebrale" title="Corteccia cerebrale – Italian" lang="it" hreflang="it" data-title="Corteccia cerebrale" 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%99%D7%A4%D7%AA_%D7%94%D7%9E%D7%95%D7%97" title="קליפת המוח – Hebrew" lang="he" hreflang="he" data-title="קליפת המוח" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kbp mw-list-item"><a href="https://kbp.wikipedia.org/wiki/Fitenum_p%C9%94%C9%94%C9%96%C9%9B" title="Fitenum pɔɔɖɛ – Kabiye" lang="kbp" hreflang="kbp" data-title="Fitenum pɔɔɖɛ" data-language-autonym="Kabɩyɛ" data-language-local-name="Kabiye" class="interlanguage-link-target"><span>Kabɩyɛ</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9C%D0%B8_%D2%9B%D1%8B%D1%80%D1%82%D1%8B%D1%81%D1%8B" title="Ми қыртысы – Kazakh" lang="kk" hreflang="kk" data-title="Ми қыртысы" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Cortex_cerebri" title="Cortex cerebri – Latin" lang="la" hreflang="la" data-title="Cortex cerebri" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Galvos_smegen%C5%B3_%C5%BEiev%C4%97" title="Galvos smegenų žievė – Lithuanian" lang="lt" hreflang="lt" data-title="Galvos smegenų žievė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Cortex_serebral" title="Cortex serebral – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Cortex serebral" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target"><span>Lingua Franca Nova</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Agyk%C3%A9reg" title="Agykéreg – Hungarian" lang="hu" hreflang="hu" data-title="Agykéreg" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Korteks_serebrum" title="Korteks serebrum – Malay" lang="ms" hreflang="ms" data-title="Korteks serebrum" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Hersenschors" title="Hersenschors – Dutch" lang="nl" hreflang="nl" data-title="Hersenschors" 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%A4%A7%E8%84%B3%E7%9A%AE%E8%B3%AA" title="大脳皮質 – Japanese" lang="ja" hreflang="ja" data-title="大脳皮質" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Hjernebark" title="Hjernebark – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Hjernebark" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Kora_m%C3%B3zgu" title="Kora mózgu – Polish" lang="pl" hreflang="pl" data-title="Kora mózgu" 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/C%C3%B3rtex_cerebral" title="Córtex cerebral – Portuguese" lang="pt" hreflang="pt" data-title="Córtex cerebral" 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/Cortex_cerebral" title="Cortex cerebral – Romanian" lang="ro" hreflang="ro" data-title="Cortex cerebral" 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%9A%D0%BE%D1%80%D0%B0_%D0%B1%D0%BE%D0%BB%D1%8C%D1%88%D0%B8%D1%85_%D0%BF%D0%BE%D0%BB%D1%83%D1%88%D0%B0%D1%80%D0%B8%D0%B9" title="Кора больших полушарий – Russian" lang="ru" hreflang="ru" data-title="Кора больших полушарий" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cerebral_cortex" title="Cerebral cortex – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cerebral cortex" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Mozgov%C3%A1_k%C3%B4ra" title="Mozgová kôra – Slovak" lang="sk" hreflang="sk" data-title="Mozgová kôra" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mo%C5%BEganska_skorja" title="Možganska skorja – Slovenian" lang="sl" hreflang="sl" data-title="Možganska skorja" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AA%D9%88%DB%8E%DA%A9%DA%B5%DB%8C_%D9%85%DB%8E%D8%B4%DA%A9" title="توێکڵی مێشک – Central Kurdish" lang="ckb" hreflang="ckb" data-title="توێکڵی مێشک" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Cerebralni_korteks" title="Cerebralni korteks – Serbian" lang="sr" hreflang="sr" data-title="Cerebralni korteks" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Cerebralni_korteks" title="Cerebralni korteks – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Cerebralni korteks" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Aivokuori" title="Aivokuori – Finnish" lang="fi" hreflang="fi" data-title="Aivokuori" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Hj%C3%A4rnbark" title="Hjärnbark – Swedish" lang="sv" hreflang="sv" data-title="Hjärnbark" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%80%E0%B8%9B%E0%B8%A5%E0%B8%B7%E0%B8%AD%E0%B8%81%E0%B8%AA%E0%B8%A1%E0%B8%AD%E0%B8%87" title="เปลือกสมอง – Thai" lang="th" hreflang="th" data-title="เปลือกสมอง" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Serebral_korteks" title="Serebral korteks – Turkish" lang="tr" hreflang="tr" data-title="Serebral korteks" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9A%D0%BE%D1%80%D0%B0_%D0%B3%D0%BE%D0%BB%D0%BE%D0%B2%D0%BD%D0%BE%D0%B3%D0%BE_%D0%BC%D0%BE%D0%B7%D0%BA%D1%83" title="Кора головного мозку – Ukrainian" lang="uk" hreflang="uk" data-title="Кора головного мозку" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ug mw-list-item"><a href="https://ug.wikipedia.org/wiki/%DA%86%D9%88%DA%AD_%D9%85%DB%90%DA%AD%DB%95_%D9%BE%D9%88%D8%B3%D8%AA%D9%84%D8%A7%D9%82_%D9%85%D8%A7%D8%AF%D8%AF%D9%89%D8%B3%D9%89" title="چوڭ مېڭە پوستلاق ماددىسى – Uyghur" lang="ug" hreflang="ug" data-title="چوڭ مېڭە پوستلاق ماددىسى" data-language-autonym="ئۇيغۇرچە / Uyghurche" data-language-local-name="Uyghur" class="interlanguage-link-target"><span>ئۇيغۇرچە / Uyghurche</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a 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</div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <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">Outer layer of the cerebrum of the mammalian 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">For the scientific journal, see <a href="/wiki/Cerebral_Cortex_(journal)" title="Cerebral Cortex (journal)">Cerebral Cortex (journal)</a>. For the cerebellar cortex, see <a href="/wiki/Cerebellum#Gross_anatomy" title="Cerebellum">Cerebellum § Gross anatomy</a>.</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:lemonchiffon">Cerebral cortex</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Cerebral_cortex,_side_view.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Cerebral_cortex%2C_side_view.svg/250px-Cerebral_cortex%2C_side_view.svg.png" decoding="async" width="250" height="151" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Cerebral_cortex%2C_side_view.svg/375px-Cerebral_cortex%2C_side_view.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Cerebral_cortex%2C_side_view.svg/500px-Cerebral_cortex%2C_side_view.svg.png 2x" data-file-width="2395" data-file-height="1449" /></a></span><div class="infobox-caption">The <a href="/wiki/Sulcus_(neuroanatomy)" title="Sulcus (neuroanatomy)">sulci</a> and <a href="/wiki/Gyrus" title="Gyrus">gyri</a> (folds and ridges) of the cortex</div></td></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Blausen_0103_Brain_Sensory%26Motor.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Blausen_0103_Brain_Sensory%26Motor.png/250px-Blausen_0103_Brain_Sensory%26Motor.png" decoding="async" width="250" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Blausen_0103_Brain_Sensory%26Motor.png/375px-Blausen_0103_Brain_Sensory%26Motor.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Blausen_0103_Brain_Sensory%26Motor.png/500px-Blausen_0103_Brain_Sensory%26Motor.png 2x" data-file-width="1600" data-file-height="960" /></a></span><div class="infobox-caption">Motor and sensory areas of the cerebral cortex</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/Cerebrum" title="Cerebrum">Cerebrum</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/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>cortex cerebri</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D002540">D002540</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=39">39</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_1494">birnlex_1494</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/14.1.09.003%20Entity%20TA98%20EN.htm">A14.1.09.003</a><br /> <a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/TA98%20Tree/Entity%20TA98%20EN/14.1.09.301%20Entity%20TA98%20EN.htm">A14.1.09.301</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=5527">5527</a>, <a rel="nofollow" class="external text" href="https://ta2viewer.openanatomy.org/?id=5528">5528</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%2Ffma61830">61830</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/Q75839" class="extiw" title="d:Q75839">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>The <b>cerebral <a href="/wiki/Cortex_(anatomy)" title="Cortex (anatomy)">cortex</a></b>, also known as the <b>cerebral mantle</b>,<sup id="cite_ref-free_1-0" class="reference"><a href="#cite_note-free-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> is the outer layer of <a href="/wiki/Neural_tissue" class="mw-redirect" title="Neural tissue">neural tissue</a> of the <a href="/wiki/Cerebrum" title="Cerebrum">cerebrum</a> of the <a href="/wiki/Brain" title="Brain">brain</a> in <a href="/wiki/Humans" class="mw-redirect" title="Humans">humans</a> and other <a href="/wiki/Mammal" title="Mammal">mammals</a>. It is the largest site of <a href="/wiki/Neuron" title="Neuron">neural integration</a> in the <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a>,<sup id="cite_ref-Saladin_2-0" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> and plays a key role in <a href="/wiki/Attention" title="Attention">attention</a>, <a href="/wiki/Perception" title="Perception">perception</a>, <a href="/wiki/Awareness" title="Awareness">awareness</a>, <a href="/wiki/Thought" title="Thought">thought</a>, <a href="/wiki/Memory" title="Memory">memory</a>, <a href="/wiki/Language" title="Language">language</a>, and <a href="/wiki/Consciousness" title="Consciousness">consciousness</a>. The cerebral cortex is the part of the brain responsible for <a href="/wiki/Cognition" title="Cognition">cognition</a>. </p><p>The six-layered <a href="/wiki/Neocortex" title="Neocortex">neocortex</a> makes up approximately 90% of the <a href="/wiki/Cortex_(anatomy)" title="Cortex (anatomy)">cortex</a>, with the <a href="/wiki/Allocortex" title="Allocortex">allocortex</a> making up the remainder.<sup id="cite_ref-Strominger_3-0" class="reference"><a href="#cite_note-Strominger-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The cortex is divided into left and right parts by the <a href="/wiki/Longitudinal_fissure" title="Longitudinal fissure">longitudinal fissure</a>, which separates the two <a href="/wiki/Cerebral_hemisphere" title="Cerebral hemisphere">cerebral hemispheres</a> that are joined beneath the cortex by the <a href="/wiki/Corpus_callosum" title="Corpus callosum">corpus callosum</a>. In most mammals, apart from small mammals that have small brains, the cerebral cortex is folded, providing a greater surface area in the confined volume of the <a href="/wiki/Neurocranium" title="Neurocranium">cranium</a>. Apart from minimising brain and cranial volume, <a href="/wiki/Gyrification" title="Gyrification">cortical folding</a> is crucial for the <a href="/wiki/Neural_circuit" title="Neural circuit">brain circuitry</a> and its functional organisation.<sup id="cite_ref-Shipp_2007_4-0" class="reference"><a href="#cite_note-Shipp_2007-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In mammals with small brains, there is no folding and the cortex is smooth.<sup id="cite_ref-embo_5-0" class="reference"><a href="#cite_note-embo-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ReferenceA_6-0" class="reference"><a href="#cite_note-ReferenceA-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>A fold or ridge in the cortex is termed a <a href="/wiki/Gyrus" title="Gyrus">gyrus</a> (plural gyri) and a groove is termed a <a href="/wiki/Sulcus_(neuroanatomy)" title="Sulcus (neuroanatomy)">sulcus</a> (plural sulci). These surface convolutions appear during <a href="/wiki/Prenatal_development" title="Prenatal development">fetal development</a> and continue to mature after birth through the process of <a href="/wiki/Gyrification" title="Gyrification">gyrification</a>. In the <a href="/wiki/Human_brain" title="Human brain">human brain</a>, the majority of the cerebral cortex is not visible from the outside, but buried in the sulci.<sup id="cite_ref-Principles_of_Neural_Science_7-0" class="reference"><a href="#cite_note-Principles_of_Neural_Science-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The major sulci and gyri mark the divisions of the cerebrum into the <a href="/wiki/Lobes_of_the_brain" title="Lobes of the brain">lobes of the brain</a>. The four major lobes are the <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal</a>, <a href="/wiki/Parietal_lobe" title="Parietal lobe">parietal</a>, <a href="/wiki/Occipital_lobe" title="Occipital lobe">occipital</a> and <a href="/wiki/Temporal_lobe" title="Temporal lobe">temporal</a> lobes. Other lobes are the <a href="/wiki/Limbic_lobe" title="Limbic lobe">limbic lobe</a>, and the <a href="/wiki/Insular_cortex" title="Insular cortex">insular cortex</a> often referred to as the <i>insular lobe</i>. </p><p>There are between 14 and 16 billion <a href="/wiki/Neuron" title="Neuron">neurons</a> in the human cerebral cortex.<sup id="cite_ref-Saladin_2-1" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These are organised into horizontal cortical layers, and radially into <a href="/wiki/Cortical_column" title="Cortical column">cortical columns</a> and <a href="/wiki/Minicolumn" class="mw-redirect" title="Minicolumn">minicolumns</a>. Cortical areas have specific functions such as movement in the <a href="/wiki/Motor_cortex" title="Motor cortex">motor cortex</a>, and sight in the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>. The motor cortex is primarily located in the <a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">precentral gyrus</a>, and the visual cortex is located in the occipital lobe. </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=Cerebral_cortex&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_motor_cortex.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Human_motor_cortex.jpg/220px-Human_motor_cortex.jpg" decoding="async" width="220" height="202" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Human_motor_cortex.jpg/330px-Human_motor_cortex.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/3/33/Human_motor_cortex.jpg 2x" data-file-width="364" data-file-height="334" /></a><figcaption>Lateral view of cerebrum showing several cortices</figcaption></figure> <p>The cerebral cortex is the outer covering of the surfaces of the cerebral hemispheres and is folded into peaks called <a href="/wiki/Gyrus" title="Gyrus">gyri</a>, and grooves called <a href="/wiki/Sulcus_(neuroanatomy)" title="Sulcus (neuroanatomy)">sulci</a>. In the <a href="/wiki/Human_brain" title="Human brain">human brain</a>, it is between 2 and 3-4 mm. thick,<sup id="cite_ref-Roberts_8-0" class="reference"><a href="#cite_note-Roberts-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> and makes up 40% of the brain's mass.<sup id="cite_ref-Saladin_2-2" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> 90% of the cerebral cortex is the six-layered <a href="/wiki/Neocortex" title="Neocortex">neocortex</a> whilst the other 10% is made up of the three/four-layered <a href="/wiki/Allocortex" title="Allocortex">allocortex</a>.<sup id="cite_ref-Saladin_2-3" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> There are between 14 and 16 billion neurons in the cortex.<sup id="cite_ref-Saladin_2-4" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These cortical neurons are organized radially in <a href="/wiki/Cortical_column" title="Cortical column">cortical columns</a>, and <a href="/wiki/Cortical_minicolumn" title="Cortical minicolumn">minicolumns</a>, in the horizontally organized layers of the cortex.<sup id="cite_ref-Lodato_9-0" class="reference"><a href="#cite_note-Lodato-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Ansen-Wilson_10-0" class="reference"><a href="#cite_note-Ansen-Wilson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>The neocortex is separable into different regions of cortex known in the plural as cortices, and include the <a href="/wiki/Motor_cortex" title="Motor cortex">motor cortex</a> and <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>. About two thirds of the cortical surface is buried in the sulci and the <a href="/wiki/Insular_cortex" title="Insular cortex">insular cortex</a> is completely hidden. The cortex is thickest over the top of a gyrus and thinnest at the bottom of a sulcus.<sup id="cite_ref-Carpenter_11-0" class="reference"><a href="#cite_note-Carpenter-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Folds">Folds</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=2" title="Edit section: Folds"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Gyrification" title="Gyrification">Gyrification</a></div> <p>The cerebral cortex is folded in a way that allows a large surface area of <a href="/wiki/Nervous_tissue" title="Nervous tissue">neural tissue</a> to fit within the confines of the <a href="/wiki/Neurocranium" title="Neurocranium">neurocranium</a>. When unfolded in the human, each <a href="/wiki/Cerebral_hemisphere" title="Cerebral hemisphere">hemispheric</a> cortex has a total surface area of about 0.12 square metres (1.3 sq ft).<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The folding is inward away from the surface of the brain, and is also present on the medial surface of each hemisphere within the <a href="/wiki/Longitudinal_fissure" title="Longitudinal fissure">longitudinal fissure</a>. Most mammals have a cerebral cortex that is convoluted with the peaks known as gyri and the troughs or grooves known as sulci. Some small mammals including some small <a href="/wiki/Rodent" title="Rodent">rodents</a> have smooth cerebral surfaces without <a href="/wiki/Gyrification" title="Gyrification">gyrification</a>.<sup id="cite_ref-ReferenceA_6-1" class="reference"><a href="#cite_note-ReferenceA-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Lobes">Lobes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=3" title="Edit section: Lobes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Lobes_of_the_brain" title="Lobes of the brain">Lobes of the brain</a></div> <p>The larger sulci and gyri mark the divisions of the cortex of the cerebrum into the <a href="/wiki/Lobes_of_the_brain" title="Lobes of the brain">lobes of the brain</a>.<sup id="cite_ref-Roberts_8-1" class="reference"><a href="#cite_note-Roberts-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> There are four main lobes: the <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal lobe</a>, <a href="/wiki/Parietal_lobe" title="Parietal lobe">parietal lobe</a>, <a href="/wiki/Temporal_lobe" title="Temporal lobe">temporal lobe</a>, and <a href="/wiki/Occipital_lobe" title="Occipital lobe">occipital lobe</a>. The <a href="/wiki/Insular_cortex" title="Insular cortex">insular cortex</a> is often included as the insular lobe.<sup id="cite_ref-Nieuwenhuys_13-0" class="reference"><a href="#cite_note-Nieuwenhuys-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Limbic_lobe" title="Limbic lobe">limbic lobe</a> is a rim of cortex on the medial side of each hemisphere and is also often included.<sup id="cite_ref-Wiley_14-0" class="reference"><a href="#cite_note-Wiley-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> There are also three lobules of the brain described: the <a href="/wiki/Paracentral_lobule" title="Paracentral lobule">paracentral lobule</a>, the <a href="/wiki/Superior_parietal_lobule" title="Superior parietal lobule">superior parietal lobule</a>, and the <a href="/wiki/Inferior_parietal_lobule" title="Inferior parietal lobule">inferior parietal lobule</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Thickness">Thickness</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=4" title="Edit section: Thickness"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For species of mammals, larger brains (in absolute terms, not just in relation to body size) tend to have thicker cortices.<sup id="cite_ref-CNSVert_15-0" class="reference"><a href="#cite_note-CNSVert-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> The smallest mammals, such as <a href="/wiki/Shrew" title="Shrew">shrews</a>, have a neocortical thickness of about 0.5 mm; the ones with the largest brains, such as humans and fin whales, have thicknesses of 2–4 mm.<sup id="cite_ref-Saladin_2-5" class="reference"><a href="#cite_note-Saladin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Roberts_8-2" class="reference"><a href="#cite_note-Roberts-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> There is an approximately <a href="/wiki/Logarithm" title="Logarithm">logarithmic</a> relationship between brain weight and cortical thickness.<sup id="cite_ref-CNSVert_15-1" class="reference"><a href="#cite_note-CNSVert-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Magnetic_resonance_imaging_of_the_brain" title="Magnetic resonance imaging of the brain">Magnetic resonance imaging of the brain</a> (MRI) makes it possible to get a measure for the thickness of the human cerebral cortex and relate it to other measures. The thickness of different cortical areas varies but in general, sensory cortex is thinner than motor cortex.<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> One study has found some positive association between the cortical thickness and <a href="/wiki/Intelligence" title="Intelligence">intelligence</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Another study has found that the <a href="/wiki/Somatosensory_cortex" class="mw-redirect" title="Somatosensory cortex">somatosensory cortex</a> is thicker in <a href="/wiki/Migraine" title="Migraine">migraine</a> patients, though it is not known if this is the result of migraine attacks, the cause of them or if both are the result of a shared cause.<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><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> A later study using a larger patient population reports no change in the cortical thickness in patients with migraine.<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> A genetic disorder of the cerebral cortex, whereby decreased folding in certain areas results in a <a href="/wiki/Microgyrus" title="Microgyrus">microgyrus</a>, where there are four layers instead of six, is in some instances seen to be related to <a href="/wiki/Dyslexia" title="Dyslexia">dyslexia</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Layers_of_neocortex"><span class="anchor" id="Layers"></span><span class="anchor" id="Laminar_pattern"></span>Layers of neocortex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=5" title="Edit section: Layers of neocortex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Minute_structure_of_the_cerebral_cortex.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Minute_structure_of_the_cerebral_cortex.jpg/220px-Minute_structure_of_the_cerebral_cortex.jpg" decoding="async" width="220" height="366" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Minute_structure_of_the_cerebral_cortex.jpg/330px-Minute_structure_of_the_cerebral_cortex.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/93/Minute_structure_of_the_cerebral_cortex.jpg/440px-Minute_structure_of_the_cerebral_cortex.jpg 2x" data-file-width="1280" data-file-height="2132" /></a><figcaption>Diagram of layers pattern. Cells grouped on left, axonal layers on right.</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Cajal_cortex_drawings.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Cajal_cortex_drawings.png/220px-Cajal_cortex_drawings.png" decoding="async" width="220" height="254" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Cajal_cortex_drawings.png/330px-Cajal_cortex_drawings.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Cajal_cortex_drawings.png/440px-Cajal_cortex_drawings.png 2x" data-file-width="672" data-file-height="777" /></a><figcaption>Three drawings of cortical lamination by <a href="/wiki/Santiago_Ramon_y_Cajal" class="mw-redirect" title="Santiago Ramon y Cajal">Santiago Ramon y Cajal</a>, each showing a vertical cross-section, with the surface of the cortex at the top. Left: <a href="/wiki/Nissl_stain" class="mw-redirect" title="Nissl stain">Nissl</a>-stained visual cortex of a human adult. Middle: Nissl-stained motor cortex of a human adult. Right: <a href="/wiki/Golgi_stain" class="mw-redirect" title="Golgi stain">Golgi</a>-stained cortex of a <style data-mw-deduplicate="TemplateStyles:r1154941027">.mw-parser-output .frac{white-space:nowrap}.mw-parser-output .frac .num,.mw-parser-output .frac .den{font-size:80%;line-height:0;vertical-align:super}.mw-parser-output .frac .den{vertical-align:sub}.mw-parser-output .sr-only{border:0;clip:rect(0,0,0,0);clip-path:polygon(0px 0px,0px 0px,0px 0px);height:1px;margin:-1px;overflow:hidden;padding:0;position:absolute;width:1px}</style><span class="frac">1<span class="sr-only">+</span><span class="num">1</span>⁄<span class="den">2</span></span> month-old infant. The Nissl stain shows the cell bodies of neurons; the Golgi stain shows the <a href="/wiki/Dendrite" title="Dendrite">dendrites</a> and axons of a random subset of neurons.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Visual_cortex_-_low_mag.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Visual_cortex_-_low_mag.jpg/220px-Visual_cortex_-_low_mag.jpg" decoding="async" width="220" height="330" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Visual_cortex_-_low_mag.jpg/330px-Visual_cortex_-_low_mag.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Visual_cortex_-_low_mag.jpg/440px-Visual_cortex_-_low_mag.jpg 2x" data-file-width="2848" data-file-height="4272" /></a><figcaption><a href="/wiki/Micrograph" title="Micrograph">Micrograph</a> showing the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a> (predominantly pink). Subcortical <a href="/wiki/White_matter" title="White matter">white matter</a> (predominantly blue) is seen at the bottom of the image. <a href="/wiki/LFB_stain" class="mw-redirect" title="LFB stain">HE-LFB stain</a>.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:NeuronGolgi.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/88/NeuronGolgi.png/220px-NeuronGolgi.png" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/88/NeuronGolgi.png/330px-NeuronGolgi.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/88/NeuronGolgi.png/440px-NeuronGolgi.png 2x" data-file-width="1400" data-file-height="1050" /></a><figcaption><a href="/wiki/Golgi%27s_method" title="Golgi's method">Golgi-stained</a> neurons in the cortex (<a href="/wiki/Macaque" title="Macaque">macaque</a>)</figcaption></figure> <p>The <a href="/wiki/Neocortex" title="Neocortex">neocortex</a> is formed of six layers, numbered I to VI, from the outermost layer I – near to the <a href="/wiki/Pia_mater" title="Pia mater">pia mater</a>, to the innermost layer VI – near to the underlying <a href="/wiki/White_matter" title="White matter">white matter</a>. Each cortical layer has a characteristic distribution of different neurons and their connections with other cortical and subcortical regions. There are direct connections between different cortical areas and indirect connections via the thalamus. </p><p>One of the clearest examples of <a href="/wiki/Laminar_organization" title="Laminar organization">cortical layering</a> is the <a href="/wiki/Line_of_Gennari" title="Line of Gennari">line of Gennari</a> in the <a href="/wiki/Visual_cortex#Primary_visual_cortex_(V1)" title="Visual cortex">primary visual cortex</a>. This is a band of whiter tissue that can be observed with the naked eye in the <a href="/wiki/Calcarine_sulcus" title="Calcarine sulcus">calcarine sulcus</a> of the occipital lobe. The line of Gennari is composed of <a href="/wiki/Axon" title="Axon">axons</a> bringing visual information from the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> into layer IV of the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>. </p><p><a href="/wiki/Staining" title="Staining">Staining</a> cross-sections of the cortex to reveal the position of neuronal cell bodies and the intracortical axon tracts allowed neuroanatomists in the early 20th century to produce a detailed description of the <i>laminar structure of the cortex</i> in different species. The work of <a href="/wiki/Korbinian_Brodmann" title="Korbinian Brodmann">Korbinian Brodmann</a> (1909) established that the mammalian neocortex is consistently divided into six layers. </p> <div class="mw-heading mw-heading4"><h4 id="Layer_I">Layer I</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=6" title="Edit section: Layer I"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Core-matrix_theory_of_thalamus" title="Core-matrix theory of thalamus">Core-matrix theory of thalamus</a></div> <p>Layer I is the <b>molecular layer</b>, and contains few scattered neurons, including <a href="/wiki/GABAergic" title="GABAergic">GABAergic</a> <a href="/wiki/Rosehip_neuron" title="Rosehip neuron">rosehip neurons</a>.<sup id="cite_ref-Allen_22-0" class="reference"><a href="#cite_note-Allen-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Layer I consists largely of extensions of apical <a href="/wiki/Dendrite" title="Dendrite">dendritic</a> tufts of <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal neurons</a> and horizontally oriented axons, as well as <a href="/wiki/Glial_cells" class="mw-redirect" title="Glial cells">glial cells</a>.<sup id="cite_ref-Shipp_2007_4-1" class="reference"><a href="#cite_note-Shipp_2007-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> During development, <a href="/wiki/Cajal%E2%80%93Retzius_cell" title="Cajal–Retzius cell">Cajal–Retzius cells</a><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> and subpial granular layer cells<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> are present in this layer. Also, some spiny <a href="/wiki/Stellate_cells" class="mw-redirect" title="Stellate cells">stellate cells</a> can be found here. Inputs to the apical tufts are thought to be crucial for the <i>feedback</i> interactions in the cerebral cortex involved in associative learning and attention.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>While it was once thought that the input to layer I came from the cortex itself,<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> it is now known that layer I across the cerebral cortex receives substantial input from <i>matrix</i> or M-type thalamus cells,<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> as opposed to <i>core</i> or C-type that go to layer IV.<sup id="cite_ref-Jones_28-0" class="reference"><a href="#cite_note-Jones-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>It is thought that layer I serves as a central hub for collecting and processing widespread information. It integrates ascending sensory inputs with top-down expectations, regulating how sensory perceptions align with anticipated outcomes. Further, layer I sorts, directs, and combines excitatory inputs, integrating them with neuromodulatory signals. Inhibitory interneurons, both within layer I and from other cortical layers, gate these signals. Together, these interactions dynamically calibrate information flow throughout the neocortex, shaping perceptions and experiences.<sup id="cite_ref-s309_29-0" class="reference"><a href="#cite_note-s309-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Layer_II">Layer II</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=7" title="Edit section: Layer II"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Layer II, the <b><a href="/wiki/External_granular_layer_(cerebral_cortex)" title="External granular layer (cerebral cortex)">external granular layer</a></b>, contains small <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal neurons</a> and numerous stellate neurons. </p> <div class="mw-heading mw-heading4"><h4 id="Layer_III">Layer III</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=8" title="Edit section: Layer III"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Layer III, the <b>external pyramidal layer</b>, contains predominantly small and medium-size pyramidal neurons, as well as non-pyramidal neurons with vertically oriented intracortical axons; layers I through III are the main target of <a href="/wiki/Commissural_fiber" title="Commissural fiber">commissural</a> corticocortical <a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">afferents</a>, and layer III is the principal source of corticocortical <a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">efferents</a>. </p> <div class="mw-heading mw-heading4"><h4 id="Layer_IV">Layer IV</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=9" title="Edit section: Layer IV"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Layer IV, the <b><a href="/wiki/Internal_granular_layer" class="mw-redirect" title="Internal granular layer">internal granular layer</a></b>, contains different types of <a href="/wiki/Stellate_cell" title="Stellate cell">stellate</a> and pyramidal cells, and is the main target of <a href="/wiki/Thalamocortical_radiations" title="Thalamocortical radiations">thalamocortical afferents</a> from thalamus type C neurons (core-type)<sup id="cite_ref-Jones_28-1" class="reference"><a href="#cite_note-Jones-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> as well as intra-hemispheric corticocortical afferents. The layers above layer IV are also referred to as supragranular layers (layers I-III), whereas the layers below are referred to as infragranular layers (layers V and VI). <a href="/wiki/African_elephant" title="African elephant">African elephants</a>, <a href="/wiki/Cetacea" title="Cetacea">cetaceans</a>, and <a href="/wiki/Hippopotamus" title="Hippopotamus">hippopotamus</a> do not have a layer IV with axons which would terminate there going instead to the inner part of layer III.<sup id="cite_ref-u035_30-0" class="reference"><a href="#cite_note-u035-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Layer_V">Layer V</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=10" title="Edit section: Layer V"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Layer V, the <b>internal pyramidal layer</b>, contains large pyramidal neurons. Axons from these leave the cortex and connect with subcortical structures including the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>. In the primary motor cortex of the frontal lobe, layer V contains giant pyramidal cells called <a href="/wiki/Betz_cell" title="Betz cell">Betz cells</a>, whose axons travel through the <a href="/wiki/Internal_capsule" title="Internal capsule">internal capsule</a>, the <a href="/wiki/Brain_stem" class="mw-redirect" title="Brain stem">brain stem</a>, and the spinal cord forming the <a href="/wiki/Corticospinal_tract" title="Corticospinal tract">corticospinal tract</a>, which is the main pathway for voluntary motor control. </p> <div class="mw-heading mw-heading4"><h4 id="Layer_VI">Layer VI</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=11" title="Edit section: Layer VI"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Layer VI, the <b>polymorphic layer</b> or <b>multiform layer</b>, contains few large pyramidal neurons and many small spindle-like pyramidal and multiform neurons; layer VI sends <a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">efferent fibers</a> to the thalamus, establishing a very precise reciprocal interconnection between the cortex and the thalamus.<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> That is, layer VI neurons from one cortical column connect with thalamus neurons that provide input to the same cortical column. These connections are both excitatory and inhibitory. Neurons send <a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">excitatory</a> fibers to neurons in the thalamus and also send collaterals to the <a href="/wiki/Thalamic_reticular_nucleus" title="Thalamic reticular nucleus">thalamic reticular nucleus</a> that <a href="/wiki/Inhibitory_postsynaptic_potential" title="Inhibitory postsynaptic potential">inhibit</a> these same thalamus neurons or ones adjacent to them.<sup id="cite_ref-Lam_32-0" class="reference"><a href="#cite_note-Lam-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> One theory is that because the inhibitory output is reduced by <a href="/wiki/Cholinergic" title="Cholinergic">cholinergic</a> input to the cerebral cortex, this provides the <a href="/wiki/Brainstem" title="Brainstem">brainstem</a> with adjustable "gain control for the relay of <a href="/wiki/Posterior_column-medial_lemniscus_pathway" class="mw-redirect" title="Posterior column-medial lemniscus pathway">lemniscal</a> inputs".<sup id="cite_ref-Lam_32-1" class="reference"><a href="#cite_note-Lam-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Columns">Columns</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=12" title="Edit section: Columns"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cortical layers are not simply stacked one over the other; there exist characteristic connections between different layers and neuronal types, which span all the thickness of the cortex. These cortical microcircuits are grouped into <a href="/wiki/Cortical_column" title="Cortical column">cortical columns</a> and <a href="/wiki/Cortical_minicolumn" title="Cortical minicolumn">minicolumns</a>.<sup id="cite_ref-Suzuki_33-0" class="reference"><a href="#cite_note-Suzuki-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> It has been proposed that the minicolumns are the basic functional units of the cortex.<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> In 1957, <a href="/wiki/Vernon_Benjamin_Mountcastle" title="Vernon Benjamin Mountcastle">Vernon Mountcastle</a> showed that the functional properties of the cortex change abruptly between laterally adjacent points; however, they are continuous in the direction perpendicular to the surface. Later works have provided evidence of the presence of functionally distinct cortical columns in the visual cortex (Hubel and <a href="/wiki/Torsten_Wiesel" title="Torsten Wiesel">Wiesel</a>, 1959),<sup id="cite_ref-pmid14403679_35-0" class="reference"><a href="#cite_note-pmid14403679-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> auditory cortex, and associative cortex. </p><p>Cortical areas that lack a layer IV are called <a href="/wiki/Agranular_cortex" title="Agranular cortex">agranular</a>. Cortical areas that have only a rudimentary layer IV are called dysgranular.<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> Information processing within each layer is determined by different temporal dynamics with that in layers II/III having a slow 2 <a href="/wiki/Hertz" title="Hertz">Hz</a> <a href="/wiki/Neural_oscillation" title="Neural oscillation">oscillation</a> while that in layer V has a fast 10–15 Hz oscillation.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Types_of_cortex">Types of cortex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=13" title="Edit section: Types of cortex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Based on the differences in <a href="/wiki/Laminar_organization" title="Laminar organization">laminar organization</a> the cerebral cortex can be classified into two types, the large area of <a href="/wiki/Neocortex" title="Neocortex">neocortex</a> which has six cell layers, and the much smaller area of <a href="/wiki/Allocortex" title="Allocortex">allocortex</a> that has three or four layers:<sup id="cite_ref-Strominger_3-1" class="reference"><a href="#cite_note-Strominger-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <ul><li>The neocortex is also known as the isocortex or neopallium and is the part of the mature cerebral cortex with six distinct layers. Examples of neocortical areas include the granular <a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">primary motor cortex</a>, and the striate <a href="/wiki/Primary_visual_cortex" class="mw-redirect" title="Primary visual cortex">primary visual cortex</a>. The neocortex has two subtypes, the <i>true isocortex</i> and the <a href="/wiki/Proisocortex" title="Proisocortex">proisocortex</a> which is a transitional region between the isocortex and the regions of the periallocortex.</li> <li>The allocortex is the part of the cerebral cortex with three or four layers, and has three subtypes, the <a href="/wiki/Paleocortex" title="Paleocortex">paleocortex</a> with three cortical laminae, the <a href="/wiki/Archicortex" title="Archicortex">archicortex</a> which has four or five, and a transitional area adjacent to the allocortex, the <a href="/wiki/Periallocortex" title="Periallocortex">periallocortex</a>. Examples of allocortex are the <a href="/wiki/Olfactory_cortex" class="mw-redirect" title="Olfactory cortex">olfactory cortex</a> and the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>.</li></ul> <p>There is a transitional area between the neocortex and the allocortex called the <a href="/wiki/Paralimbic_cortex" title="Paralimbic cortex">paralimbic cortex</a>, where layers 2, 3 and 4 are merged. This area incorporates the proisocortex of the neocortex and the periallocortex of the allocortex. In addition, the cerebral cortex may be classified into four <a href="/wiki/Lobes_of_the_brain" title="Lobes of the brain">lobes</a>: the <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal lobe</a>, <a href="/wiki/Temporal_lobe" title="Temporal lobe">temporal lobe</a>, the <a href="/wiki/Parietal_lobe" title="Parietal lobe">parietal lobe</a>, and the <a href="/wiki/Occipital_lobe" title="Occipital lobe">occipital lobe</a>, named from their overlying bones of the skull. </p> <div class="mw-heading mw-heading2"><h2 id="Blood_supply_and_drainage">Blood supply and drainage</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=14" title="Edit section: Blood supply and drainage"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cerebral_circulation" title="Cerebral circulation">Cerebral circulation</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cerebrovascular_System.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Cerebrovascular_System.png/220px-Cerebrovascular_System.png" decoding="async" width="220" height="205" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Cerebrovascular_System.png/330px-Cerebrovascular_System.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Cerebrovascular_System.png/440px-Cerebrovascular_System.png 2x" data-file-width="603" data-file-height="561" /></a><figcaption>Arterial supply showing the regions supplied by the posterior, middle, and anterior <a href="/wiki/Cerebral_arteries" title="Cerebral arteries">cerebral arteries</a>.</figcaption></figure> <p>Blood supply to the cerebral cortex is part of the <a href="/wiki/Cerebral_circulation" title="Cerebral circulation">cerebral circulation</a>. <a href="/wiki/Cerebral_arteries" title="Cerebral arteries">Cerebral arteries</a> supply the blood that <a href="/wiki/Perfusion" title="Perfusion">perfuses</a> the cerebrum. This arterial blood carries oxygen, glucose, and other nutrients to the cortex. <a href="/wiki/Cerebral_veins" title="Cerebral veins">Cerebral veins</a> drain the deoxygenated blood, and metabolic wastes including carbon dioxide, back to the heart. </p><p>The main arteries supplying the cortex are the <a href="/wiki/Anterior_cerebral_artery" title="Anterior cerebral artery">anterior cerebral artery</a>, the <a href="/wiki/Middle_cerebral_artery" title="Middle cerebral artery">middle cerebral artery</a>, and the <a href="/wiki/Posterior_cerebral_artery" title="Posterior cerebral artery">posterior cerebral artery</a>. The anterior cerebral artery supplies the anterior portions of the brain, including most of the frontal lobe. The middle cerebral artery supplies the parietal lobes, temporal lobes, and parts of the occipital lobes. The middle cerebral artery splits into two branches to supply the left and right hemisphere, where they branch further. The posterior cerebral artery supplies the occipital lobes. </p><p>The <a href="/wiki/Circle_of_Willis" title="Circle of Willis">circle of Willis</a> is the main blood system that deals with blood supply in the cerebrum and cerebral cortex. </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Cerebral_vascular_territories.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Cerebral_vascular_territories.jpg/220px-Cerebral_vascular_territories.jpg" decoding="async" width="220" height="215" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Cerebral_vascular_territories.jpg/330px-Cerebral_vascular_territories.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Cerebral_vascular_territories.jpg/440px-Cerebral_vascular_territories.jpg 2x" data-file-width="1024" data-file-height="999" /></a><figcaption>Cortical blood supply</figcaption></figure> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Development">Development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=15" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Development_of_the_cerebral_cortex" title="Development of the cerebral cortex">Development of the cerebral cortex</a></div> <p>The <a href="/wiki/Prenatal_development" title="Prenatal development">prenatal development</a> of the cerebral cortex is a complex and finely tuned process called <a href="/wiki/Development_of_the_human_cerebral_cortex" class="mw-redirect" title="Development of the human cerebral cortex">corticogenesis</a>, influenced by the interplay between genes and the environment.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neural_tube">Neural tube</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=16" title="Edit section: Neural tube"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cerebral cortex develops from the most anterior part, the forebrain region, of the <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>.<sup id="cite_ref-Wolpert_39-0" class="reference"><a href="#cite_note-Wolpert-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid10498281_40-0" class="reference"><a href="#cite_note-pmid10498281-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Neural_plate" title="Neural plate">neural plate</a> folds and closes to form the <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>. From the cavity inside the neural tube develops the <a href="/wiki/Ventricular_system" title="Ventricular system">ventricular system</a>, and, from the <a href="/wiki/Neuroepithelial_cell" title="Neuroepithelial cell">neuroepithelial cells</a> of its walls, the <a href="/wiki/Neuron" title="Neuron">neurons</a> and <a href="/wiki/Neuroglia" class="mw-redirect" title="Neuroglia">glia</a> of the nervous system. The most anterior (front, or cranial) part of the neural plate, the <a href="/wiki/Prosencephalon" class="mw-redirect" title="Prosencephalon">prosencephalon</a>, which is evident before <a href="/wiki/Neurulation" title="Neurulation">neurulation</a> begins, gives rise to the cerebral hemispheres and later cortex.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Cortical_neuron_development">Cortical neuron development</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=17" title="Edit section: Cortical neuron development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a> and <a href="/wiki/Neuroepithelial_cell" title="Neuroepithelial cell">Neuroepithelial cell</a></div> <p>Cortical neurons are generated within the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a>, next to the <a href="/wiki/Ventricular_system" title="Ventricular system">ventricles</a>. At first, this zone contains <a href="/wiki/Neural_stem_cell" title="Neural stem cell">neural stem cells</a>, that transition to <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glial cells</a>–progenitor cells, which divide to produce glial cells and neurons.<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> </p> <div class="mw-heading mw-heading4"><h4 id="Radial_glia">Radial glia</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=18" title="Edit section: Radial glia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg/220px-Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg" decoding="async" width="220" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg/330px-Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg/440px-Neurogenesis_and_Differentiation_of_Cortical_Layers.jpg 2x" data-file-width="747" data-file-height="409" /></a><figcaption>Neurogenesis is shown in red and lamination is shown in blue. Adapted from (Sur et al. 2001)</figcaption></figure> <p>The cerebral cortex is composed of a heterogenous population of cells that give rise to different cell types. The majority of these cells are derived from <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glia</a> migration that form the different cell types of the neocortex and it is a period associated with an increase in <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a>. Similarly, the process of neurogenesis regulates lamination to form the different layers of the cortex. During this process there is an increase in the restriction of cell fate that begins with earlier <a href="/wiki/Progenitor_cell" title="Progenitor cell">progenitors</a> giving rise to any cell type in the cortex and later progenitors giving rise only to <a href="/wiki/Neuron" title="Neuron">neurons</a> of superficial layers. This differential cell fate creates an inside-out topography in the cortex with younger neurons in superficial layers and older neurons in deeper layers. In addition, laminar neurons are stopped in <a href="/wiki/Cell_cycle" title="Cell cycle">S</a> or <a href="/wiki/G2_phase" title="G2 phase">G2 phase</a> in order to give a fine distinction between the different cortical layers. Laminar differentiation is not fully complete until after birth since during development laminar neurons are still sensitive to extrinsic signals and environmental cues.<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> </p><p>Although the majority of the cells that compose the cortex are derived locally from radial glia there is a subset population of neurons that <a href="/wiki/Cell_migration" title="Cell migration">migrate</a> from other regions. Radial glia give rise to neurons that are pyramidal in shape and use <a href="/wiki/Glutamate_(neurotransmitter)" title="Glutamate (neurotransmitter)">glutamate</a> as a <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a>, however these migrating cells contribute neurons that are stellate-shaped and use <a href="/wiki/Gamma-Aminobutyric_acid" class="mw-redirect" title="Gamma-Aminobutyric acid">GABA</a> as their main neurotransmitter. These GABAergic neurons are generated by progenitor cells in the <a href="/wiki/Medial_ganglionic_eminence" class="mw-redirect" title="Medial ganglionic eminence">medial ganglionic eminence</a> (MGE) that migrate tangentially to the cortex via the <a href="/wiki/Subventricular_zone" title="Subventricular zone">subventricular zone</a>. This migration of GABAergic neurons is particularly important since <a href="/wiki/GABA_receptor" title="GABA receptor">GABA receptors</a> are excitatory during development. This excitation is primarily driven by the flux of chloride ions through the GABA receptor, however in adults chloride concentrations shift causing an inward flux of chloride that <a href="/wiki/Hyperpolarization_(biology)" title="Hyperpolarization (biology)">hyperpolarizes</a> <a href="/wiki/Postsynaptic_neuron" class="mw-redirect" title="Postsynaptic neuron">postsynaptic neurons</a>.<sup id="cite_ref-Sanes_2012_44-0" class="reference"><a href="#cite_note-Sanes_2012-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> The glial fibers produced in the first divisions of the progenitor cells are radially oriented, spanning the thickness of the cortex from the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a> to the outer, <a href="/wiki/Pia_mater" title="Pia mater">pial</a> surface, and provide scaffolding for the migration of neurons outwards from the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a>.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><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><p>At birth there are very few <a href="/wiki/Dendrite" title="Dendrite">dendrites</a> present on the cortical neuron's cell body, and the axon is undeveloped. During the first year of life the dendrites become dramatically increased in number, such that they can accommodate up to a hundred thousand <a href="/wiki/Synapse" title="Synapse">synaptic connections</a> with other neurons. The axon can develop to extend a long way from the cell body.<sup id="cite_ref-Gilbert2_47-0" class="reference"><a href="#cite_note-Gilbert2-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Asymmetric_division">Asymmetric division</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=19" title="Edit section: Asymmetric division"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first divisions of the progenitor cells are symmetric, which duplicates the total number of progenitor cells at each <a href="/wiki/Mitosis" title="Mitosis">mitotic cycle</a>. Then, some progenitor cells begin to divide asymmetrically, producing one postmitotic cell that migrates along the radial glial fibers, leaving the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a>, and one progenitor cell, which continues to divide until the end of development, when it differentiates into a <a href="/wiki/Astrocyte" title="Astrocyte">glial cell</a> or an <a href="/wiki/Ependyma" title="Ependyma">ependymal cell</a>. As the <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a> of <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> is elongated, in what is seen as selective cell-cycle lengthening, the newly born neurons migrate to more superficial layers of the cortex.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> The migrating daughter cells become the <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal cells</a> of the cerebral cortex.<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> The development process is time ordered and regulated by hundreds of genes and <a href="/wiki/Epigenetic_Regulation_of_Neurogenesis" class="mw-redirect" title="Epigenetic Regulation of Neurogenesis">epigenetic regulatory mechanisms</a>.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Layer_organization">Layer organization</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=20" title="Edit section: Layer organization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Human_Cortical_Development.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Human_Cortical_Development.png/220px-Human_Cortical_Development.png" decoding="async" width="220" height="254" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Human_Cortical_Development.png/330px-Human_Cortical_Development.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Human_Cortical_Development.png/440px-Human_Cortical_Development.png 2x" data-file-width="2303" data-file-height="2656" /></a><figcaption>Human cortical development between 26 and 39 week gestational age</figcaption></figure> <p>The <a href="/wiki/Laminar_organization" title="Laminar organization">layered structure</a> of the mature cerebral cortex is formed during development. The first pyramidal neurons generated migrate out of the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a> and <a href="/wiki/Subventricular_zone" title="Subventricular zone">subventricular zone</a>, together with <a href="/wiki/Reelin" title="Reelin">reelin</a>-producing <a href="/wiki/Cajal%E2%80%93Retzius_cell" title="Cajal–Retzius cell">Cajal–Retzius neurons</a>, from the <b>preplate</b>. Next, a cohort of neurons migrating into the middle of the preplate divides this transient layer into the superficial <b>marginal zone</b>, which will become layer I of the mature neocortex, and the <a href="/wiki/Subplate" title="Subplate">subplate</a>,<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> forming a middle layer called the <b>cortical plate</b>. These cells will form the deep layers of the mature cortex, layers five and six. Later born neurons migrate radially into the cortical plate past the deep layer neurons, and become the upper layers (two to four). Thus, the layers of the cortex are created in an inside-out order.<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> The only exception to this inside-out sequence of <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a> occurs in the layer I of <a href="/wiki/Primate" title="Primate">primates</a>, in which, in contrast to <a href="/wiki/Rodent" title="Rodent">rodents</a>, neurogenesis continues throughout the entire period of <a href="/wiki/Corticogenesis" class="mw-redirect" title="Corticogenesis">corticogenesis</a>.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cortical_patterning">Cortical patterning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=21" title="Edit section: Cortical patterning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Emx2_and_Pax6_Expression.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Emx2_and_Pax6_Expression.png/220px-Emx2_and_Pax6_Expression.png" decoding="async" width="220" height="155" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Emx2_and_Pax6_Expression.png/330px-Emx2_and_Pax6_Expression.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/30/Emx2_and_Pax6_Expression.png/440px-Emx2_and_Pax6_Expression.png 2x" data-file-width="1352" data-file-height="951" /></a><figcaption>Depicted in blue, Emx2 is highly expressed at the caudomedial pole and dissipates outward. Pax6 expression is represented in purple and is highly expressed at the rostral lateral pole. (Adapted from Sanes, D., Reh, T., & Harris, W. (2012). <i>Development of the Nervous System</i> (3rd ed.). Burlington: Elsevier Science)</figcaption></figure> <p>The map of functional cortical areas, which include primary motor and visual cortex, originates from a '<a href="/wiki/Protomap_(neuroscience)" title="Protomap (neuroscience)">protomap</a>',<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> which is regulated by molecular signals such as <a href="/wiki/Fibroblast_growth_factor" title="Fibroblast growth factor">fibroblast growth factor</a> <a href="/wiki/FGF8" class="mw-redirect" title="FGF8">FGF8</a> early in embryonic development.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> These signals regulate the size, shape, and position of cortical areas on the surface of the cortical primordium, in part by regulating gradients of <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factor</a> expression, through a process called <a href="/wiki/Cortical_patterning" title="Cortical patterning">cortical patterning</a>. Examples of such transcription factors include the genes <a href="/wiki/EMX2" title="EMX2">EMX2</a> and <a href="/wiki/PAX6" title="PAX6">PAX6</a>.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Together, both <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> form an opposing gradient of expression. <a href="/wiki/PAX6" title="PAX6">Pax6</a> is highly expressed at the <a href="/wiki/Anatomical_terms_of_location" title="Anatomical terms of location">rostral lateral</a> pole, while <a href="/wiki/EMX2" title="EMX2">Emx2</a> is highly expressed in the <a href="/wiki/Anatomical_terms_of_location" title="Anatomical terms of location">caudomedial</a> pole. The establishment of this gradient is important for proper development. For example, <a href="/wiki/Mutation" title="Mutation">mutations</a> in Pax6 can cause expression levels of Emx2 to expand out of its normal expression domain, which would ultimately lead to an expansion of the areas normally derived from the caudal medial cortex, such as the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>. On the contrary, if mutations in Emx2 occur, it can cause the Pax6-expressing domain to expand and result in the <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal</a> and <a href="/wiki/Motor_cortex" title="Motor cortex">motor cortical</a> regions enlarging. Therefore, researchers believe that similar gradients and <a href="/wiki/Cell_signaling" title="Cell signaling">signaling centers</a> next to the cortex could contribute to the regional expression of these transcription factors.<sup id="cite_ref-Sanes_2012_44-1" class="reference"><a href="#cite_note-Sanes_2012-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Two very well studied patterning signals for the cortex include <a href="/wiki/Fibroblast_growth_factor" title="Fibroblast growth factor">FGF</a> and <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a>. If FGFs are <a href="/wiki/Protein_production" title="Protein production">misexpressed</a> in different areas of the developing cortex, <a href="/wiki/Cortical_patterning" title="Cortical patterning">cortical patterning</a> is disrupted. Specifically, when <a href="/wiki/FGF8" class="mw-redirect" title="FGF8">Fgf8</a> is increased in the <a href="/wiki/Anatomical_terms_of_location" title="Anatomical terms of location">anterior</a> pole, Emx2 is <a href="/wiki/Downregulation_and_upregulation" title="Downregulation and upregulation">downregulated</a> and a <a href="/wiki/Caudal_(anatomical_term)" class="mw-redirect" title="Caudal (anatomical term)">caudal</a> shift in the cortical region occurs. This ultimately causes an expansion of the rostral regions. Therefore, Fgf8 and other FGFs play a role in the regulation of expression of Emx2 and Pax6 and represent how the cerebral cortex can become specialized for different functions.<sup id="cite_ref-Sanes_2012_44-2" class="reference"><a href="#cite_note-Sanes_2012-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Rapid expansion of the cortical surface area is regulated by the amount of self-renewal of <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glial cells</a> and is partly regulated by <a href="/wiki/Fibroblast_growth_factor" title="Fibroblast growth factor">FGF</a> and <a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch genes</a>.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> During the period of cortical neurogenesis and layer formation, many higher mammals begin the process of <a href="/wiki/Gyrification" title="Gyrification">gyrification</a>, which generates the characteristic folds of the cerebral cortex.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Gyrification is regulated by a DNA-associated protein <a href="/wiki/Trnp1" class="mw-redirect" title="Trnp1">Trnp1</a><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> and by FGF and <a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">SHH</a> signaling.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=22" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Pallium_(neuroanatomy)#Evolution" title="Pallium (neuroanatomy)">Pallium (neuroanatomy) § Evolution</a></div> <p>Of all the different brain regions, the cerebral cortex shows the largest evolutionary variation and has evolved most recently.<sup id="cite_ref-ReferenceA_6-2" class="reference"><a href="#cite_note-ReferenceA-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In contrast to the highly conserved circuitry of the <a href="/wiki/Medulla_oblongata" title="Medulla oblongata">medulla oblongata</a>, for example, which serves critical functions such as regulation of heart and respiration rates, many areas of the cerebral cortex are not strictly necessary for survival. Thus, the evolution of the cerebral cortex has seen the advent and modification of new functional areas—particularly association areas that do not directly receive input from outside the cortex.<sup id="cite_ref-ReferenceA_6-3" class="reference"><a href="#cite_note-ReferenceA-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>A key theory of cortical evolution is embodied in the <a href="/wiki/Radial_unit_hypothesis" title="Radial unit hypothesis">radial unit hypothesis</a> and related <a href="/wiki/Protomap_(neuroscience)" title="Protomap (neuroscience)">protomap</a> hypothesis, first proposed by Rakic.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> This theory states that new cortical areas are formed by the addition of new radial units, which is accomplished at the <a href="/wiki/Stem_cell" title="Stem cell">stem cell</a> level. The protomap hypothesis states that the cellular and molecular identity and characteristics of neurons in each cortical area are specified by cortical <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a>, known as <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glial cells</a>, in a primordial map. This map is controlled by secreted signaling <a href="/wiki/Protein" title="Protein">proteins</a> and downstream <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a>.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </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=Cerebral_cortex&action=edit&section=23" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Constudproc.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Constudproc.png/220px-Constudproc.png" decoding="async" width="220" height="249" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Constudproc.png/330px-Constudproc.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Constudproc.png/440px-Constudproc.png 2x" data-file-width="763" data-file-height="865" /></a><figcaption>Some functional areas of cortex</figcaption></figure> <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=Cerebral_cortex&action=edit&section=24" title="Edit section: Connections"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cerebral cortex is connected to various subcortical structures such as the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> and the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>, sending information to them along <a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">efferent connections</a> and receiving information from them via <a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">afferent connections</a>. Most sensory information is routed to the cerebral cortex via the thalamus. Olfactory information, however, passes through the <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a> to the olfactory cortex (<a href="/wiki/Piriform_cortex" title="Piriform cortex">piriform cortex</a>). The majority of connections are from one area of the cortex to another, rather than from subcortical areas; <a href="/wiki/Valentino_Braitenberg" title="Valentino Braitenberg">Braitenberg</a> and Schüz (1998) claim that in primary sensory areas, at the cortical level where the input fibers terminate, up to 20% of the synapses are supplied by extracortical afferents but that in other areas and other layers the percentage is likely to be much lower.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cortical_areas">Cortical areas</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=25" title="Edit section: Cortical areas"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Cortical_map" title="Cortical map">Cortical map</a></div> <p>The whole of the cerebral cortex was divided into 52 different areas in an early presentation by <a href="/wiki/Korbinian_Brodmann" title="Korbinian Brodmann">Korbinian Brodmann</a>. These areas, known as <a href="/wiki/Brodmann_area" title="Brodmann area">Brodmann areas</a>, are based on their <a href="/wiki/Cytoarchitecture" title="Cytoarchitecture">cytoarchitecture</a> but also relate to various functions. An example is Brodmann area 17, which is the <a href="/wiki/Visual_cortex#Primary_visual_cortex_(V1)" title="Visual cortex">primary visual cortex</a>. </p><p>In more general terms the cortex is typically described as comprising three parts: sensory, motor, and association areas. </p> <div class="mw-heading mw-heading4"><h4 id="Sensory_areas">Sensory areas</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=26" title="Edit section: Sensory areas"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Blausen_0102_Brain_Motor%26Sensory.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Blausen_0102_Brain_Motor%26Sensory.png/220px-Blausen_0102_Brain_Motor%26Sensory.png" decoding="async" width="220" height="205" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Blausen_0102_Brain_Motor%26Sensory.png/330px-Blausen_0102_Brain_Motor%26Sensory.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Blausen_0102_Brain_Motor%26Sensory.png/440px-Blausen_0102_Brain_Motor%26Sensory.png 2x" data-file-width="1600" data-file-height="1493" /></a><figcaption>Motor and sensory regions of the cerebral cortex</figcaption></figure> <p>The sensory areas are the cortical areas that receive and process information from the <a href="/wiki/Senses" class="mw-redirect" title="Senses">senses</a>. Parts of the cortex that receive sensory inputs from the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> are called primary sensory areas. The senses of vision, hearing, and touch are served by the primary visual cortex, primary <a href="/wiki/Auditory_cortex" title="Auditory cortex">auditory cortex</a> and <a href="/wiki/Primary_somatosensory_cortex" title="Primary somatosensory cortex">primary somatosensory cortex</a> respectively. In general, the two hemispheres receive information from the opposite (contralateral) side of the <a href="/wiki/Human_body" title="Human body">body</a>. For example, the right primary somatosensory cortex receives information from the left limbs, and the right visual cortex receives information from the left visual <a href="/wiki/Receptive_field" title="Receptive field">field</a>. </p><p>The organization of sensory maps in the cortex reflects that of the corresponding sensing organ, in what is known as a <a href="/wiki/Topographic_map_(Neuroanatomy)" class="mw-redirect" title="Topographic map (Neuroanatomy)">topographic map</a>. Neighboring points in the primary <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a>, for example, correspond to neighboring points in the <a href="/wiki/Retina" title="Retina">retina</a>. This topographic map is called a <a href="/wiki/Retinotopy" title="Retinotopy">retinotopic map</a>. In the same way, there exists a <a href="/wiki/Tonotopy" title="Tonotopy">tonotopic map</a> in the primary auditory cortex and a <a href="/wiki/Somatotopy" class="mw-redirect" title="Somatotopy">somatotopic map</a> in the primary sensory cortex. This last topographic map of the body onto the <a href="/wiki/Posterior_central_gyrus" class="mw-redirect" title="Posterior central gyrus">posterior central gyrus</a> has been illustrated as a deformed human representation, the somatosensory <a href="/wiki/Cortical_homunculus" title="Cortical homunculus">homunculus</a>, where the size of different body parts reflects the relative density of their innervation. Areas with much sensory innervation, such as the fingertips and the lips, require more cortical area to process finer sensation. </p> <div class="mw-heading mw-heading4"><h4 id="Motor_areas">Motor areas</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=27" title="Edit section: Motor areas"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The motor areas are located in both hemispheres of the cortex. The motor areas are very closely related to the control of voluntary movements, especially fine fragmented movements performed by the hand. The right half of the motor area controls the left side of the body, and vice versa. </p><p>Two areas of the cortex are commonly referred to as motor: </p> <ul><li><a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a>, which <i>executes</i> voluntary movements <sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor areas</a> and <a href="/wiki/Premotor_cortex" title="Premotor cortex">premotor cortex</a>, which <i>select</i> voluntary movements.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup></li></ul> <p>In addition, motor functions have been described for: </p> <ul><li><a href="/wiki/Posterior_parietal_cortex" title="Posterior parietal cortex">Posterior parietal cortex</a>, which guides voluntary movements in space</li> <li><a href="/wiki/Dorsolateral_prefrontal_cortex" title="Dorsolateral prefrontal cortex">Dorsolateral prefrontal cortex</a>, which decides which voluntary movements to make according to higher-order instructions, rules, and self-generated thoughts.</li></ul> <p>Just underneath the cerebral cortex are interconnected subcortical masses of grey matter called <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> (or nuclei). The basal ganglia receive input from the substantia nigra of the midbrain and motor areas of the cerebral cortex, and send signals back to both of these locations. They are involved in motor control. They are found lateral to the thalamus. The main components of the basal ganglia are the <a href="/wiki/Caudate_nucleus" title="Caudate nucleus">caudate nucleus</a>, the <a href="/wiki/Putamen" title="Putamen">putamen</a>, the <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a>, the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a>, the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a>, and the <a href="/wiki/Subthalamic_nucleus" title="Subthalamic nucleus">subthalamic nucleus</a>. The putamen and globus pallidus are also collectively known as the <a href="/wiki/Lentiform_nucleus" title="Lentiform nucleus">lentiform nucleus</a>, because together they form a lens-shaped body. The putamen and caudate nucleus are also collectively called the <a href="/wiki/Corpus_striatum" class="mw-redirect" title="Corpus striatum">corpus striatum</a> after their striped appearance.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Association_areas">Association areas</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=28" title="Edit section: Association areas"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg/220px-Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg" decoding="async" width="220" height="131" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg/330px-Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/22/Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg/440px-Cortical_areas_that_have_been_shown_to_be_involved_in_speech_processing_fnhum-06-00099-g005.jpg 2x" data-file-width="545" data-file-height="325" /></a><figcaption>Cortical areas involved in speech processing.</figcaption></figure> <p>The association areas are the parts of the cerebral cortex that do not belong to the primary regions. They function to produce a meaningful <a href="/wiki/Perception" title="Perception">perceptual experience</a> of the world, enable us to interact effectively, and support abstract thinking and language. The <a href="/wiki/Parietal_lobe" title="Parietal lobe">parietal</a>, <a href="/wiki/Temporal_lobe" title="Temporal lobe">temporal</a>, and <a href="/wiki/Occipital_lobe" title="Occipital lobe">occipital lobes</a> – all located in the posterior part of the cortex – integrate sensory information and information stored in memory. The <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal lobe</a> or prefrontal association complex is involved in planning actions and movement, as well as abstract thought. Globally, the association areas are organized as distributed networks.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Each network connects areas distributed across widely spaced regions of the cortex. Distinct networks are positioned adjacent to one another yielding a complex series of interwoven networks. The specific organization of the association networks is debated with evidence for interactions, hierarchical relationships, and competition between networks. </p><p>In humans, association networks are particularly important to language function. In the past it was theorized that language abilities are localized in <a href="/wiki/Broca%27s_area" title="Broca's area">Broca's area</a> in areas of the left <a href="/wiki/Inferior_frontal_gyrus" title="Inferior frontal gyrus">inferior frontal gyrus</a>, <a href="/wiki/Brodmann_area_44" title="Brodmann area 44">BA44</a> and <a href="/wiki/Brodmann_area_45" title="Brodmann area 45">BA45</a>, for language expression and in <a href="/wiki/Wernicke%27s_area" title="Wernicke's area">Wernicke's area</a> <a href="/wiki/Brodmann_area" title="Brodmann area">BA22</a>, for language reception. However, the processes of language expression and reception have been shown to occur in areas other than just those structures around the <a href="/wiki/Lateral_sulcus" title="Lateral sulcus">lateral sulcus</a>, including the frontal lobe, <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>, <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>, and <a href="/wiki/Pons" title="Pons">pons</a>.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> </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=Cerebral_cortex&action=edit&section=29" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Central_nervous_system_disease" title="Central nervous system disease">Central nervous system disease</a> and <a href="/wiki/Developmental_toxicity" title="Developmental toxicity">Developmental toxicity</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Middle_Cerebral_Artery_occlusion._Kentar_et_al_Acta_Neuroch_2020.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4d/Middle_Cerebral_Artery_occlusion._Kentar_et_al_Acta_Neuroch_2020.gif/507px-Middle_Cerebral_Artery_occlusion._Kentar_et_al_Acta_Neuroch_2020.gif" decoding="async" width="507" height="364" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/4/4d/Middle_Cerebral_Artery_occlusion._Kentar_et_al_Acta_Neuroch_2020.gif 1.5x" data-file-width="632" data-file-height="454" /></a><figcaption>Hemodynamic changes observed on gyrencephalic brain cortex after an arterial vessel occlusion in IOS. The video has a speed of 50x to better appreciate the <a href="/wiki/Cortical_spreading_depression" title="Cortical spreading depression">spreading depolarization</a> over the brain cortex. Pictures are dynamically subtracted to a reference picture 40 s before. First we see the initial area of change at the exact moment where the middle cerebral artery group (left) is occluded. The area is highlighted with a white line. Later we appreciate the signal produced by Spreading Depolarizations. We see markedly the front of waves.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> https://doi.org/10.1007/s00701-019-04132-8</figcaption></figure> <p><a href="/wiki/Neurodegenerative_disease" title="Neurodegenerative disease">Neurodegenerative diseases</a> such as <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a>, show as a marker, an atrophy of the grey matter of the cerebral cortex.<sup id="cite_ref-Nakazawa_76-0" class="reference"><a href="#cite_note-Nakazawa-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p>Other <a href="/wiki/Central_nervous_system_disease" title="Central nervous system disease">diseases of the central nervous system</a> include <a href="/wiki/Neurological_disorder" title="Neurological disorder">neurological disorders</a> such as <a href="/wiki/Epilepsy" title="Epilepsy">epilepsy</a>, <a href="/wiki/Movement_disorder" title="Movement disorder">movement disorders</a>, and <a href="/wiki/Aphasia#Boston_classification" title="Aphasia">different types of aphasia</a> (difficulties in speech expression or comprehension). </p><p><a href="/wiki/Brain_damage" class="mw-redirect" title="Brain damage">Brain damage</a> from disease or trauma, can involve damage to a specific lobe such as in <a href="/wiki/Frontal_lobe_disorder" title="Frontal lobe disorder">frontal lobe disorder</a>, and associated functions will be affected. The <a href="/wiki/Blood%E2%80%93brain_barrier" title="Blood–brain barrier">blood–brain barrier</a> that serves to protect the brain from infection can become compromised allowing entry to <a href="/wiki/Pathogen" title="Pathogen">pathogens</a>. </p><p>The <a href="/wiki/Prenatal_development" title="Prenatal development">developing fetus</a> is susceptible to a range of environmental factors that can cause <a href="/wiki/Birth_defect" title="Birth defect">birth defects</a> and problems in later development. Maternal alcohol consumption for example can cause <a href="/wiki/Fetal_alcohol_spectrum_disorder" title="Fetal alcohol spectrum disorder">fetal alcohol spectrum disorder</a>.<sup id="cite_ref-Mukherjee_2006_77-0" class="reference"><a href="#cite_note-Mukherjee_2006-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> Other factors that can cause neurodevelopment disorders are <a href="/wiki/Environmental_toxicants_and_fetal_development" title="Environmental toxicants and fetal development">toxicants</a> such as <a href="/wiki/Drug" title="Drug">drugs</a>, and exposure to <a href="/wiki/Radiation" title="Radiation">radiation</a> as from <a href="/wiki/X-ray" title="X-ray">X-rays</a>. Infections can also affect the development of the cortex. A viral infection is one of the causes of <a href="/wiki/Lissencephaly" title="Lissencephaly">lissencephaly</a>, which results in a smooth cortex without <a href="/wiki/Gyrification" title="Gyrification">gyrification</a>. </p><p>A type of <a href="/wiki/Electrocorticography" title="Electrocorticography">electrocorticography</a> called <a href="/wiki/Cortical_stimulation_mapping" title="Cortical stimulation mapping">cortical stimulation mapping</a> is an invasive procedure that involves placing <a href="/wiki/Electrode" title="Electrode">electrodes</a> directly onto the exposed brain in order to localise the functions of specific areas of the cortex. It is used in clinical and therapeutic applications including pre-surgical mapping.<sup id="cite_ref-Tarapore_78-0" class="reference"><a href="#cite_note-Tarapore-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genes_associated_with_cortical_disorders">Genes associated with cortical disorders</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=30" title="Edit section: Genes associated with cortical disorders"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are a number of genetic mutations that can cause a wide range of <a href="/wiki/Genetic_disorder" title="Genetic disorder">genetic disorders</a> of the cerebral cortex, including <a href="/wiki/Microcephaly" title="Microcephaly">microcephaly</a>, <a href="/wiki/Schizencephaly" title="Schizencephaly">schizencephaly</a> and types of <a href="/wiki/Lissencephaly" title="Lissencephaly">lissencephaly</a>.<sup id="cite_ref-Walsh_79-0" class="reference"><a href="#cite_note-Walsh-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Chromosome_abnormality" title="Chromosome abnormality">Chromosome abnormalities</a> can also result causing a number of <a href="/wiki/Neurodevelopmental_disorder" title="Neurodevelopmental disorder">neurodevelopmental disorders</a> such as <a href="/wiki/Fragile_X_syndrome" title="Fragile X syndrome">fragile X syndrome</a> and <a href="/wiki/Rett_syndrome" title="Rett syndrome">Rett syndrome</a>. </p><p><a href="/wiki/MCPH1" class="mw-redirect" title="MCPH1">MCPH1</a> codes for <a href="/wiki/Microcephalin" title="Microcephalin">microcephalin</a>, and disorders in this and in <a href="/wiki/ASPM_(gene)" title="ASPM (gene)">ASPM</a> are associated with microcephaly.<sup id="cite_ref-Walsh_79-1" class="reference"><a href="#cite_note-Walsh-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> Mutations in the gene <a href="/wiki/NBS1" class="mw-redirect" title="NBS1">NBS1</a> that codes for <a href="/wiki/Nibrin" title="Nibrin">nibrin</a> can cause <a href="/wiki/Nijmegen_breakage_syndrome" title="Nijmegen breakage syndrome">Nijmegen breakage syndrome</a>, characterised by microcephaly.<sup id="cite_ref-Walsh_79-2" class="reference"><a href="#cite_note-Walsh-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p>Mutations in <a href="/wiki/EMX2" title="EMX2">EMX2</a>,<sup id="cite_ref-nih_80-0" class="reference"><a href="#cite_note-nih-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/COL4A1" class="mw-redirect" title="COL4A1">COL4A1</a> are associated with <a href="/wiki/Schizencephaly" title="Schizencephaly">schizencephaly</a>,<sup id="cite_ref-Smigiel_81-0" class="reference"><a href="#cite_note-Smigiel-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> a condition marked by the absence of large parts of the cerebral hemispheres. </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=31" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1909, <a href="/wiki/Korbinian_Brodmann" title="Korbinian Brodmann">Korbinian Brodmann</a> distinguished 52 different regions of the cerebral cortex based on their cytoarchitecture. These are known as <a href="/wiki/Brodmann_area" title="Brodmann area">Brodmann areas</a>.<sup id="cite_ref-McGraw-Hill_Medical_2013_82-0" class="reference"><a href="#cite_note-McGraw-Hill_Medical_2013-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Rafael_Lorente_de_N%C3%B3" title="Rafael Lorente de Nó">Rafael Lorente de Nó</a>, a student of <a href="/wiki/Santiago_Ramon_y_Cajal" class="mw-redirect" title="Santiago Ramon y Cajal">Santiago Ramon y Cajal</a>, identified more than 40 different types of cortical neurons based on the distribution of their dendrites and axons.<sup id="cite_ref-McGraw-Hill_Medical_2013_82-1" class="reference"><a href="#cite_note-McGraw-Hill_Medical_2013-82"><span class="cite-bracket">[</span>82<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=Cerebral_cortex&action=edit&section=32" title="Edit section: Other animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Barrel_cortex" title="Barrel cortex">Barrel cortex</a></div> <p>The cerebral cortex is derived from the <a href="/wiki/Pallium_(neuroanatomy)" title="Pallium (neuroanatomy)">pallium</a>, a layered structure found in the <a href="/wiki/Prosencephalon" class="mw-redirect" title="Prosencephalon">forebrain</a> of all <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a>. The basic form of the pallium is a cylindrical layer enclosing fluid-filled ventricles. Around the circumference of the cylinder are four zones, the dorsal pallium, medial pallium, ventral pallium, and lateral pallium, which are thought to be <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous</a> to the <a href="/wiki/Neocortex" title="Neocortex">neocortex</a>, <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, <a href="/wiki/Amygdala" title="Amygdala">amygdala</a>, and <a href="/wiki/Piriform_cortex" title="Piriform cortex">olfactory cortex</a>, respectively. </p><p>In <a href="/wiki/Avian_brain" title="Avian brain">avian brains</a>, evidence suggests the <a href="/wiki/Avian_pallium" title="Avian pallium">avian pallium</a>'s neuroarchitecture to be reminiscent of the mammalian cerebral cortex.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> The avian pallium has also been suggested to be an equivalent neural basis for <a href="/wiki/Animal_consciousness" title="Animal consciousness">consciousness</a>.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </p><p>Until recently no counterpart to the cerebral cortex had been recognized in invertebrates. However, a study published in the journal <i>Cell</i> in 2010, based on gene expression profiles, reported strong affinities between the cerebral cortex and the <a href="/wiki/Mushroom_bodies" title="Mushroom bodies">mushroom bodies</a> of the <a href="/wiki/Nereididae" title="Nereididae">ragworm</a> <i><a href="/wiki/Platynereis_dumerilii" title="Platynereis dumerilii">Platynereis dumerilii</a></i>.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> Mushroom bodies are structures in the brains of many types of worms and arthropods that are known to play important roles in learning and memory; the genetic evidence indicates a common evolutionary origin, and therefore indicates that the origins of the earliest precursors of the cerebral cortex date back to the <a href="/wiki/Precambrian" title="Precambrian">Precambrian</a> era. </p> <div class="mw-heading mw-heading2"><h2 id="Additional_images">Additional images</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cerebral_cortex&action=edit&section=33" title="Edit section: Additional images"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Lateral_surface_of_cerebral_cortex_-_gyri.png" class="mw-file-description" title="Lateral surface of the human cerebral cortex"><img alt="Lateral surface of the human cerebral cortex" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Lateral_surface_of_cerebral_cortex_-_gyri.png/120px-Lateral_surface_of_cerebral_cortex_-_gyri.png" decoding="async" width="120" height="78" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Lateral_surface_of_cerebral_cortex_-_gyri.png/180px-Lateral_surface_of_cerebral_cortex_-_gyri.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/Lateral_surface_of_cerebral_cortex_-_gyri.png/240px-Lateral_surface_of_cerebral_cortex_-_gyri.png 2x" data-file-width="1236" data-file-height="800" /></a></span></div> <div class="gallerytext">Lateral surface of the human cerebral cortex</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png" class="mw-file-description" title="Medial surface of the human cerebral cortex"><img alt="Medial surface of the human cerebral cortex" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png/120px-Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png" decoding="async" width="120" height="76" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png/180px-Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/15/Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png/240px-Medial_surface_of_cerebral_cortex_-_entorhinal_cortex.png 2x" data-file-width="1179" data-file-height="747" /></a></span></div> <div class="gallerytext">Medial surface of the human cerebral cortex</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Brainmaps-macaque-hippocampus.jpg" class="mw-file-description" title="Tissue slice from the brain of an adult macaque monkey. The cerebral cortex is depicted in dark violet."><img alt="Tissue slice from the brain of an adult macaque monkey. The cerebral cortex is depicted in dark violet." src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9a/Brainmaps-macaque-hippocampus.jpg/120px-Brainmaps-macaque-hippocampus.jpg" decoding="async" width="120" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9a/Brainmaps-macaque-hippocampus.jpg/180px-Brainmaps-macaque-hippocampus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9a/Brainmaps-macaque-hippocampus.jpg/240px-Brainmaps-macaque-hippocampus.jpg 2x" data-file-width="745" data-file-height="587" /></a></span></div> <div class="gallerytext">Tissue slice from the brain of an adult <a href="/wiki/Macaque" title="Macaque">macaque</a> monkey. The cerebral cortex is depicted in dark violet. </div> </li> </ul> <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=Cerebral_cortex&action=edit&section=34" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 30em;"> <ul><li><a href="/wiki/Brain%E2%80%93computer_interface" title="Brain–computer interface">Brain–computer interface</a></li> <li><a href="/wiki/Cortical_dysplasia" class="mw-redirect" title="Cortical dysplasia">Cortical dysplasia</a></li> <li><a href="/wiki/Cortical_homunculus" title="Cortical homunculus">Cortical homunculus</a></li> <li><a href="/wiki/Eloquent_cortex" title="Eloquent cortex">Eloquent cortex</a></li> <li><a href="/wiki/EMX1" title="EMX1">EMX1</a></li> <li><a href="/wiki/Gray_matter_heterotopia" title="Gray matter heterotopia">Gray matter heterotopia</a></li> <li><a href="/wiki/Limbic_system" title="Limbic system">Limbic system</a></li> <li><a href="/wiki/List_of_regions_in_the_human_brain" title="List of regions in the human brain">List of regions in the human brain</a></li> <li><a href="/wiki/Insular_cortex" title="Insular cortex">Insular cortex</a></li></ul> </div> <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=Cerebral_cortex&action=edit&section=35" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-free-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-free_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.thefreedictionary.com/cerebral+mantle">"cerebral mantle"</a>. <i>TheFreeDictionary.com</i><span class="reference-accessdate">. Retrieved <span class="nowrap">9 May</span> 2024</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=TheFreeDictionary.com&rft.atitle=cerebral+mantle&rft_id=https%3A%2F%2Fwww.thefreedictionary.com%2Fcerebral%2Bmantle&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACerebral+cortex" class="Z3988"></span></span> </li> <li id="cite_note-Saladin-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Saladin_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Saladin_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Saladin_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Saladin_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Saladin_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Saladin_2-5"><sup><i><b>f</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSaladin2011" class="citation book cs1">Saladin K (2011). <i>Human anatomy</i> (3rd ed.). 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class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://braininfo.rprc.washington.edu/Scripts/hiercentraldirectory.aspx?ID=20">hier-20</a> at <a href="/wiki/NeuroNames" title="NeuroNames">NeuroNames</a></li> <li><a rel="nofollow" class="external text" href="http://brainmaps.org/index.php?q=cerebral%20cortex">Stained brain slice images which include the "cerebral cortex"</a> at the <a href="/wiki/BrainMaps" title="BrainMaps">BrainMaps project</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20041229070957/http://webvision.med.utah.edu/VisualCortex.html">"The primary visual cortex"</a>, Webvision: Comprehensive article about the structure and function of the primary visual cortex.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20160722010322/http://webvision.med.utah.edu/imageswv/BasicCells.jpg">"Basic cell types"</a>, Webvision: Image of the basic cell types of the monkey cerebral cortex.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20161117212320/http://ccdb.ucsd.edu/sand/main?stype=lite&keyword=cerebral%20cortex&Submit=Go&event=display&start=1">Cerebral Cortex – Cell Centered Database</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul 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aria-labelledby="Anatomy_of_the_cerebral_cortex_of_the_human_brain" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cerebral_cortex" title="Template:Cerebral cortex"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cerebral_cortex" title="Template talk:Cerebral cortex"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cerebral_cortex" title="Special:EditPage/Template:Cerebral cortex"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Anatomy_of_the_cerebral_cortex_of_the_human_brain" style="font-size:114%;margin:0 4em">Anatomy of the <a class="mw-selflink selflink">cerebral cortex</a> of the <a href="/wiki/Human_brain" title="Human brain">human brain</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Frontal_lobe" title="Frontal lobe">Frontal lobe</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%">Superolateral</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">Prefrontal</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/Superior_frontal_gyrus" title="Superior frontal gyrus">Superior frontal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_4" title="Brodmann area 4">4</a></li> <li><a href="/wiki/Brodmann_area_6" title="Brodmann area 6">6</a></li> <li><a href="/wiki/Brodmann_area_8" title="Brodmann area 8">8</a></li></ul></li> <li><a href="/wiki/Middle_frontal_gyrus" title="Middle frontal gyrus">Middle frontal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_9" title="Brodmann area 9">9</a></li> <li><a href="/wiki/Brodmann_area_10" title="Brodmann area 10">10</a></li> <li><a href="/wiki/Brodmann_area_46" title="Brodmann area 46">46</a></li></ul></li></ul> <ul><li><a href="/wiki/Inferior_frontal_gyrus" title="Inferior frontal gyrus">Inferior frontal gyrus</a>: <a href="/wiki/Brodmann_area_11" title="Brodmann area 11">11</a></li> <li><a href="/wiki/Brodmann_area_47" title="Brodmann area 47">47</a>-<a href="/wiki/Orbital_part_of_inferior_frontal_gyrus" title="Orbital part of inferior frontal gyrus">Pars orbitalis</a></li> <li><a href="/wiki/Broca%27s_area" title="Broca's area">Broca's area</a> <ul><li><a href="/wiki/Brodmann_area_44" title="Brodmann area 44">44</a>-<a href="/wiki/Opercular_part_of_inferior_frontal_gyrus" class="mw-redirect" title="Opercular part of inferior frontal gyrus">Pars opercularis</a></li> <li><a href="/wiki/Brodmann_area_45" title="Brodmann area 45">45</a>-<a href="/wiki/Triangular_part_of_inferior_frontal_gyrus" class="mw-redirect" title="Triangular part of inferior frontal gyrus">Pars triangularis</a></li></ul></li></ul> <ul><li><a href="/wiki/Superior_frontal_sulcus" title="Superior frontal sulcus">Superior frontal sulcus</a></li> <li><a href="/wiki/Inferior_frontal_sulcus" title="Inferior frontal sulcus">Inferior frontal sulcus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Precentral_gyrus" title="Precentral gyrus">Precentral</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/Precentral_gyrus" title="Precentral gyrus">Precentral gyrus</a></li> <li><a href="/wiki/Precentral_sulcus" title="Precentral sulcus">Precentral sulcus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/inferior</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">Prefrontal</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/Superior_frontal_gyrus" title="Superior frontal gyrus">Superior frontal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_4" title="Brodmann area 4">4</a></li> <li><a href="/wiki/Brodmann_area_6" title="Brodmann area 6">6</a></li></ul></li> <li><a href="/wiki/Medial_frontal_gyrus" title="Medial frontal gyrus">Medial frontal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_8" title="Brodmann area 8">8</a></li> <li><a href="/wiki/Brodmann_area_9" title="Brodmann area 9">9</a></li></ul></li></ul> <ul><li><a href="/wiki/Paraterminal_gyrus" class="mw-redirect" title="Paraterminal gyrus">Paraterminal gyrus</a>/<a href="/wiki/Paraolfactory_area" class="mw-redirect" title="Paraolfactory area">Paraolfactory area</a> <ul><li><a href="/wiki/Brodmann_area_12" title="Brodmann area 12">12</a></li></ul></li> <li><a href="/wiki/Straight_gyrus" title="Straight gyrus">Straight gyrus</a> <ul><li><a href="/wiki/Brodmann_area_11" title="Brodmann area 11">11</a></li></ul></li> <li><a href="/wiki/Orbital_gyri" title="Orbital gyri">Orbital gyri</a>/<a href="/wiki/Orbitofrontal_cortex" title="Orbitofrontal cortex">Orbitofrontal cortex</a> <ul><li><a href="/wiki/Brodmann_area_10" title="Brodmann area 10">10</a></li> <li><a href="/wiki/Brodmann_area_11" title="Brodmann area 11">11</a></li> <li><a href="/wiki/Brodmann_area_12" title="Brodmann area 12">12</a></li></ul></li> <li><a href="/wiki/Ventromedial_prefrontal_cortex" title="Ventromedial prefrontal cortex">Ventromedial prefrontal cortex</a> <ul><li><a href="/wiki/Brodmann_area_10" title="Brodmann area 10">10</a></li></ul></li> <li><a href="/wiki/Subcallosal_area" title="Subcallosal area">Subcallosal area</a> <ul><li><a href="/wiki/Brodmann_area_25" title="Brodmann area 25">25</a></li></ul></li></ul> <ul><li><a href="/wiki/Olfactory_sulcus" class="mw-redirect" title="Olfactory sulcus">Olfactory sulcus</a></li> <li><a href="/wiki/Orbital_sulcus" title="Orbital sulcus">Orbital sulcus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Precentral_gyrus" title="Precentral gyrus">Precentral</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/Paracentral_lobule" title="Paracentral lobule">Paracentral lobule</a> <ul><li><a href="/wiki/Brodmann_area_4" title="Brodmann area 4">4</a></li></ul></li> <li><a href="/wiki/Paracentral_sulcus" title="Paracentral sulcus">Paracentral sulcus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Both</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a> <ul><li><a href="/wiki/Brodmann_area_4" title="Brodmann area 4">4</a></li></ul></li> <li><a href="/wiki/Premotor_cortex" title="Premotor cortex">Premotor cortex</a> <ul><li><a href="/wiki/Brodmann_area_6" title="Brodmann area 6">6</a></li></ul></li> <li><a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor area</a> <ul><li><a href="/wiki/Brodmann_area_6" title="Brodmann area 6">6</a></li></ul></li> <li><a href="/wiki/Supplementary_eye_field" title="Supplementary eye field">Supplementary eye field</a> <ul><li><a href="/wiki/Brodmann_area_6" title="Brodmann area 6">6</a></li></ul></li> <li><a href="/wiki/Frontal_eye_fields" title="Frontal eye fields">Frontal eye fields</a> <ul><li><a href="/wiki/Brodmann_area_8" title="Brodmann area 8">8</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Parietal_lobe" title="Parietal lobe">Parietal lobe</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%">Superolateral</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/Superior_parietal_lobule" title="Superior parietal lobule">Superior parietal lobule</a> <ul><li><a href="/wiki/Brodmann_area_5" title="Brodmann area 5">5</a></li> <li><a href="/wiki/Brodmann_area_7" title="Brodmann area 7">7</a></li></ul></li> <li><a href="/wiki/Inferior_parietal_lobule" title="Inferior parietal lobule">Inferior parietal lobule</a> <ul><li><a href="/wiki/Brodmann_area_40" title="Brodmann area 40">40</a>-<a href="/wiki/Supramarginal_gyrus" title="Supramarginal gyrus">Supramarginal gyrus</a></li> <li><a href="/wiki/Brodmann_area_39" title="Brodmann area 39">39</a>-<a href="/wiki/Angular_gyrus" title="Angular gyrus">Angular gyrus</a></li></ul></li> <li><a href="/wiki/Parietal_operculum" class="mw-redirect" title="Parietal operculum">Parietal operculum</a> <ul><li><a href="/wiki/Brodmann_area_43" title="Brodmann area 43">43</a></li></ul></li></ul> <ul><li><a href="/wiki/Intraparietal_sulcus" title="Intraparietal sulcus">Intraparietal sulcus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/inferior</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/Paracentral_lobule" title="Paracentral lobule">Paracentral lobule</a> <ul><li><a href="/wiki/Brodmann_area_1" class="mw-redirect" title="Brodmann area 1">1</a></li> <li><a href="/wiki/Brodmann_area_2" class="mw-redirect" title="Brodmann area 2">2</a></li> <li><a href="/wiki/Brodmann_area_3" class="mw-redirect" title="Brodmann area 3">3</a></li> <li><a href="/wiki/Brodmann_area_5" title="Brodmann area 5">5</a></li></ul></li> <li><a href="/wiki/Precuneus" title="Precuneus">Precuneus</a> <ul><li><a href="/wiki/Brodmann_area_7" title="Brodmann area 7">7</a></li></ul></li> <li><a href="/wiki/Marginal_sulcus" title="Marginal sulcus">Marginal sulcus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Both</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/Postcentral_gyrus" title="Postcentral gyrus">Postcentral gyrus</a>/<a href="/wiki/Primary_somatosensory_cortex" title="Primary somatosensory cortex">Primary somatosensory cortex</a> <ul><li><a href="/wiki/Brodmann_areas_3,_1_and_2" class="mw-redirect" title="Brodmann areas 3, 1 and 2">3, 1 and 2</a></li></ul></li> <li><a href="/wiki/Secondary_somatosensory_cortex" title="Secondary somatosensory cortex">Secondary somatosensory cortex</a> <ul><li><a href="/wiki/Brodmann_area_5" title="Brodmann area 5">5</a></li></ul></li> <li><a href="/wiki/Posterior_parietal_cortex" title="Posterior parietal cortex">Posterior parietal cortex</a> <ul><li><a href="/wiki/Brodmann_area_7" title="Brodmann area 7">7</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Occipital_lobe" title="Occipital lobe">Occipital lobe</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%">Superolateral</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/Occipital_pole_of_cerebrum" class="mw-redirect" title="Occipital pole of cerebrum">Occipital pole of cerebrum</a></li> <li><a href="/wiki/Occipital_gyri" title="Occipital gyri">Occipital gyri</a></li> <li><a href="/wiki/Lateral_occipital_sulcus" title="Lateral occipital sulcus">Lateral occipital gyrus</a> <ul><li><a href="/wiki/Brodmann_area_18" title="Brodmann area 18">18</a></li> <li><a href="/wiki/Brodmann_area_19" title="Brodmann area 19">19</a></li></ul></li> <li><a href="/wiki/Lunate_sulcus" title="Lunate sulcus">Lunate sulcus</a></li> <li><a href="/wiki/Transverse_occipital_sulcus" title="Transverse occipital sulcus">Transverse occipital sulcus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/inferior</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/Visual_cortex" title="Visual cortex">Visual cortex</a> <ul><li><a href="/wiki/Brodmann_area_17" class="mw-redirect" title="Brodmann area 17">17</a></li></ul></li> <li><a href="/wiki/Cuneus" title="Cuneus">Cuneus</a></li> <li><a href="/wiki/Lingual_gyrus" title="Lingual gyrus">Lingual gyrus</a></li></ul> <ul><li><a href="/wiki/Calcarine_sulcus" title="Calcarine sulcus">Calcarine sulcus</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/Temporal_lobe" title="Temporal lobe">Temporal lobe</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%">Superolateral</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/Transverse_temporal_gyrus" title="Transverse temporal gyrus">Transverse temporal gyrus</a>/<a href="/wiki/Auditory_cortex" title="Auditory cortex">Auditory cortex</a> <ul><li><a href="/wiki/Brodmann_areas_41_and_42" title="Brodmann areas 41 and 42">41 and 42</a></li></ul></li> <li><a href="/wiki/Superior_temporal_gyrus" title="Superior temporal gyrus">Superior temporal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_38" title="Brodmann area 38">38</a></li> <li><a href="/wiki/Brodmann_area_22" title="Brodmann area 22">22</a>/<a href="/wiki/Wernicke%27s_area" title="Wernicke's area">Wernicke's area</a> (<a href="/wiki/Planum_temporale" title="Planum temporale">Planum temporale</a>)</li></ul></li> <li><a href="/wiki/Superior_temporal_sulcus" title="Superior temporal sulcus">Superior temporal sulcus</a></li> <li><a href="/wiki/Middle_temporal_gyrus" title="Middle temporal gyrus">Middle temporal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_21" title="Brodmann area 21">21</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/inferior</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/Occipitotemporal_sulcus" title="Occipitotemporal sulcus">Occipitotemporal sulcus</a></li> <li><a href="/wiki/Fusiform_gyrus" title="Fusiform gyrus">Fusiform gyrus</a> <ul><li><a href="/wiki/Brodmann_area_37" title="Brodmann area 37">37</a></li></ul></li> <li><a href="/wiki/Medial_temporal_lobe" class="mw-redirect" title="Medial temporal lobe">Medial temporal lobe</a> <ul><li><a href="/wiki/Brodmann_area_27" title="Brodmann area 27">27</a></li> <li><a href="/wiki/Brodmann_area_28" title="Brodmann area 28">28</a></li> <li><a href="/wiki/Brodmann_area_34" title="Brodmann area 34">34</a></li> <li><a href="/wiki/Brodmann_area_35" class="mw-redirect" title="Brodmann area 35">35</a></li> <li><a href="/wiki/Brodmann_area_36" class="mw-redirect" title="Brodmann area 36">36</a></li></ul></li></ul> <ul><li><a href="/wiki/Inferior_temporal_sulcus" class="mw-redirect" title="Inferior temporal sulcus">Inferior temporal sulcus</a></li> <li><a href="/wiki/Inferior_temporal_gyrus" title="Inferior temporal gyrus">Inferior temporal gyrus</a> <ul><li><a href="/wiki/Brodmann_area_20" title="Brodmann area 20">20</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sulcus_(neuroanatomy)" title="Sulcus (neuroanatomy)">Interlobar<br />sulci/fissures</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%">Superolateral</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/Central_sulcus" title="Central sulcus">Central (frontal+parietal)</a></li> <li><a href="/wiki/Lateral_sulcus" title="Lateral sulcus">Lateral (frontal+parietal+temporal)</a></li> <li><a href="/wiki/Parieto-occipital_sulcus" title="Parieto-occipital sulcus">Parieto-occipital</a></li> <li><a href="/wiki/Preoccipital_notch" title="Preoccipital notch">Preoccipital notch</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/inferior</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/Longitudinal_fissure" title="Longitudinal fissure">Longitudinal fissure</a></li> <li><a href="/wiki/Cingulate_sulcus" title="Cingulate sulcus">Cingulate (frontal+cingulate)</a></li> <li><a href="/wiki/Collateral_fissure" title="Collateral fissure">Collateral (temporal+occipital)</a></li> <li><a href="/wiki/Callosal_sulcus" class="mw-redirect" title="Callosal sulcus">Callosal sulcus</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/Limbic_lobe" title="Limbic lobe">Limbic lobe</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/Parahippocampal_gyrus" title="Parahippocampal gyrus">Parahippocampal gyrus</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><i>anterior</i> <ul><li><a href="/wiki/Entorhinal_cortex" title="Entorhinal cortex">Entorhinal cortex</a></li> <li><a href="/wiki/Perirhinal_cortex" title="Perirhinal cortex">Perirhinal cortex</a></li> <li><a href="/wiki/Postrhinal_cortex" title="Postrhinal cortex">Postrhinal cortex</a></li></ul></li> <li><a href="/wiki/Posterior_parahippocampal_gyrus" title="Posterior parahippocampal gyrus">Posterior parahippocampal gyrus</a></li> <li><a href="/wiki/Prepyriform_area" title="Prepyriform area">Prepyriform area</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cingulate_cortex" title="Cingulate cortex">Cingulate cortex</a>/<a href="/wiki/Cingulate_gyrus" class="mw-redirect" title="Cingulate gyrus">gyrus</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/Subgenual_area_25" class="mw-redirect" title="Subgenual area 25">Subgenual area</a> <ul><li><a href="/wiki/Brodmann_area_25" title="Brodmann area 25">25</a></li></ul></li> <li><a href="/wiki/Anterior_cingulate_cortex" title="Anterior cingulate cortex">Anterior cingulate</a> <ul><li><a href="/wiki/Brodmann_area_24" title="Brodmann area 24">24</a></li> <li><a href="/wiki/Brodmann_area_32" title="Brodmann area 32">32</a></li> <li><a href="/wiki/Brodmann_area_33" title="Brodmann area 33">33</a></li></ul></li> <li><a href="/wiki/Posterior_cingulate" class="mw-redirect" title="Posterior cingulate">Posterior cingulate</a> <ul><li><a href="/wiki/Brodmann_area_23" title="Brodmann area 23">23</a></li> <li><a href="/wiki/Brodmann_area_31" title="Brodmann area 31">31</a></li></ul></li></ul> <ul><li><a href="/wiki/Isthmus_of_cingulate_gyrus" title="Isthmus of cingulate gyrus">Isthmus of cingulate gyrus</a>: <a href="/wiki/Retrosplenial_region" class="mw-redirect" title="Retrosplenial region">Retrosplenial cortex</a> <ul><li><a href="/wiki/Brodmann_area_26" title="Brodmann area 26">26</a></li> <li><a href="/wiki/Brodmann_area_29" title="Brodmann area 29">29</a></li> <li><a href="/wiki/Brodmann_area_30" title="Brodmann area 30">30</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hippocampal_formation" title="Hippocampal formation">Hippocampal formation</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/Hippocampal_sulcus" title="Hippocampal sulcus">Hippocampal sulcus</a></li> <li><a href="/wiki/Fimbria_of_hippocampus" class="mw-redirect" title="Fimbria of hippocampus">Fimbria of hippocampus</a></li> <li><a href="/wiki/Dentate_gyrus" title="Dentate gyrus">Dentate gyrus</a></li> <li><a href="/wiki/Rhinal_sulcus" title="Rhinal sulcus">Rhinal sulcus</a></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/Indusium_griseum" title="Indusium griseum">Indusium griseum</a></li> <li><a href="/wiki/Uncus" title="Uncus">Uncus</a></li> <li><a href="/wiki/Amygdala" title="Amygdala">Amygdala</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/Insular_cortex" title="Insular cortex">Insular cortex</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Insular_cortex" title="Insular cortex">Insular cortex</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Operculum_(brain)" title="Operculum (brain)">Operculum</a></li> <li><a href="/wiki/Poles_of_cerebral_hemispheres" class="mw-redirect" title="Poles of cerebral hemispheres">Poles of cerebral hemispheres</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background: transparent; padding: 0px;"><div><i>Some categorizations are approximations, and some <a href="/wiki/Brodmann_area" title="Brodmann area">Brodmann areas</a> span gyri.</i></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Types_of_cerebral_cortex" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cortex_types" title="Template:Cortex types"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cortex_types" title="Template talk:Cortex types"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cortex_types" title="Special:EditPage/Template:Cortex types"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Types_of_cerebral_cortex" style="font-size:114%;margin:0 4em">Types of <a class="mw-selflink selflink">cerebral cortex</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Structure types</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Isocortex <ul><li>True isocortex or <a href="/wiki/Neocortex" title="Neocortex">Neocortex</a></li> <li><a href="/wiki/Proisocortex" title="Proisocortex">Proisocortex</a></li></ul></li> <li><a href="/wiki/Allocortex" title="Allocortex">Allocortex</a> <ul><li><a href="/wiki/Paleocortex" title="Paleocortex">Paleocortex</a></li> <li><a href="/wiki/Archicortex" title="Archicortex">Archicortex</a></li></ul></li></ul> <ul><li><a href="/wiki/Line_of_Gennari" title="Line of Gennari">Line of Gennari</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Layers</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Internal_granular_layer_(cerebral_cortex)" title="Internal granular layer (cerebral cortex)">Internal granular layer</a></li> <li><a href="/wiki/External_granular_layer_(cerebral_cortex)" title="External granular layer (cerebral cortex)">External granular layer</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cells</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Betz_cell" title="Betz cell">Betz cell</a></li> <li><a href="/wiki/Pyramidal_cell" title="Pyramidal cell">Pyramidal cell</a></li> <li><a href="/wiki/Stellate_cell" title="Stellate cell">Stellate cell</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox authority-control" 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