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href="#Firing_classifications"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Firing classifications</span> </div> </a> <ul id="toc-Firing_classifications-sublist" class="vector-toc-list"> <li id="toc-RSad" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#RSad"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.1</span> <span>RSad</span> </div> </a> <ul id="toc-RSad-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RSna" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#RSna"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.2</span> <span>RSna</span> </div> </a> <ul id="toc-RSna-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-IB" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#IB"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.3</span> <span>IB</span> </div> </a> <ul id="toc-IB-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Molecular_classifications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Molecular_classifications"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Molecular classifications</span> </div> </a> <ul id="toc-Molecular_classifications-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</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-Corticospinal_tract" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Corticospinal_tract"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Corticospinal tract</span> </div> </a> <ul id="toc-Corticospinal_tract-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cognition" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cognition"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Cognition</span> </div> </a> <ul id="toc-Cognition-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Memory_and_learning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Memory_and_learning"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Memory and learning</span> </div> </a> <ul id="toc-Memory_and_learning-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div 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">Pyramidal cell</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 23 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-23" 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">23 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AE%D9%84%D9%8A%D8%A9_%D9%87%D8%B1%D9%85%D9%8A%D8%A9" title="خلية هرمية – Arabic" lang="ar" hreflang="ar" data-title="خلية هرمية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Piramidal_h%C3%BCceyr%C9%99" title="Piramidal hüceyrə – Azerbaijani" lang="az" hreflang="az" data-title="Piramidal hüceyrə" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Neurones_piramidals" title="Neurones piramidals – Catalan" lang="ca" hreflang="ca" data-title="Neurones piramidals" 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/Pyramidov%C3%BD_neuron" title="Pyramidový neuron – Czech" lang="cs" hreflang="cs" data-title="Pyramidový neuron" 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/Pyramidecelle" title="Pyramidecelle – Danish" lang="da" hreflang="da" data-title="Pyramidecelle" 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/Pyramidenzelle" title="Pyramidenzelle – German" lang="de" hreflang="de" data-title="Pyramidenzelle" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Neurona_piramidal" title="Neurona piramidal – Spanish" lang="es" hreflang="es" data-title="Neurona piramidal" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D9%84%D9%88%D9%84%E2%80%8C%D9%87%D8%A7%DB%8C_%D9%87%D8%B1%D9%85%DB%8C" title="سلول‌های هرمی – Persian" lang="fa" hreflang="fa" data-title="سلول‌های هرمی" 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/Cellule_pyramidale" title="Cellule pyramidale – French" lang="fr" hreflang="fr" data-title="Cellule pyramidale" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/C%C3%A9lula_piramidal" title="Célula piramidal – Galician" lang="gl" hreflang="gl" data-title="Célula piramidal" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%ED%94%BC%EB%9D%BC%EB%AF%B8%EB%93%9C_%EC%84%B8%ED%8F%AC" title="피라미드 세포 – Korean" lang="ko" hreflang="ko" data-title="피라미드 세포" 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/%D4%B2%D6%80%D5%A3%D5%A1%D5%B1%D6%87_%D5%B6%D5%A5%D5%B5%D6%80%D5%B8%D5%B6" title="Բրգաձև նեյրոն – Armenian" lang="hy" hreflang="hy" data-title="Բրգաձև նեյրոն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Cellula_piramidale" title="Cellula piramidale – Italian" lang="it" hreflang="it" data-title="Cellula piramidale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Piramidinis_neuronas" title="Piramidinis neuronas – Lithuanian" lang="lt" hreflang="lt" data-title="Piramidinis neuronas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Piramidecellen" title="Piramidecellen – Dutch" lang="nl" hreflang="nl" data-title="Piramidecellen" 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/%E9%8C%90%E4%BD%93%E7%B4%B0%E8%83%9E_(%E7%A5%9E%E7%B5%8C%E7%B4%B0%E8%83%9E)" title="錐体細胞 (神経細胞) – Japanese" lang="ja" hreflang="ja" data-title="錐体細胞 (神経細胞)" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Neuron_piramidowy" title="Neuron piramidowy – Polish" lang="pl" hreflang="pl" data-title="Neuron piramidowy" 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%A9lula_piramidal" title="Célula piramidal – Portuguese" lang="pt" hreflang="pt" data-title="Célula piramidal" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%B8%D1%80%D0%B0%D0%BC%D0%B8%D0%B4%D0%B0%D0%BB%D1%8C%D0%BD%D1%8B%D0%B9_%D0%BD%D0%B5%D0%B9%D1%80%D0%BE%D0%BD" title="Пирамидальный нейрон – Russian" lang="ru" hreflang="ru" data-title="Пирамидальный нейрон" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Pyramidcell" title="Pyramidcell – Swedish" lang="sv" hreflang="sv" data-title="Pyramidcell" 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%8B%E0%B8%A5%E0%B8%A5%E0%B9%8C%E0%B8%9E%E0%B8%B5%E0%B8%A3%E0%B8%B0%E0%B8%A1%E0%B8%B4%E0%B8%94" title="เซลล์พีระมิด – Thai" lang="th" hreflang="th" data-title="เซลล์พีระมิด" 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/Piramidal_h%C3%BCcreler" title="Piramidal hücreler – Turkish" lang="tr" hreflang="tr" data-title="Piramidal hücreler" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E9%94%A5%E4%BD%93%E7%BB%86%E8%83%9E" title="锥体细胞 – Chinese" lang="zh" hreflang="zh" data-title="锥体细胞" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q2116409#sitelinks-wikipedia" title="Edit interlanguage 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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">Pyramidal cell</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:GolgiStainedPyramidalCell.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/GolgiStainedPyramidalCell.jpg/250px-GolgiStainedPyramidalCell.jpg" decoding="async" width="250" height="166" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/GolgiStainedPyramidalCell.jpg/375px-GolgiStainedPyramidalCell.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/6/6d/GolgiStainedPyramidalCell.jpg 2x" data-file-width="409" data-file-height="272" /></a></span><div class="infobox-caption">A human <a href="/wiki/Neocortex" title="Neocortex">neocortical</a> pyramidal cell stained via <a href="/wiki/Golgi%27s_method" title="Golgi's method">Golgi's method</a>. The <a href="/wiki/Apical_dendrite" title="Apical dendrite">apical dendrite</a> extends vertically above the <a href="/wiki/Soma_(biology)" title="Soma (biology)">soma</a> (cell body) and the numerous <a href="/wiki/Basal_dendrite" title="Basal dendrite">basal dendrites</a> radiate laterally from the base of the cell body.</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">Location</th><td class="infobox-data"><a href="/wiki/Cerebral_cortex#Layer_III" title="Cerebral cortex">Layer III</a> and <a href="/wiki/Cerebral_cortex#Layer_V" title="Cerebral cortex">layer V</a> of the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a>, the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, and <a href="/wiki/Amygdala" title="Amygdala">amygdala</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Shape</th><td class="infobox-data">Multipolar pyramidal</td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Function</th><td class="infobox-data">Excitatory projection neuron</td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Neurotransmitter</th><td class="infobox-data"><a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">Glutamate</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D017966">D017966</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/sao862606388">sao862606388</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Histologica" title="Terminologia Histologica">TH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/PDF/TH%20Chapter%20H1.00.pdf">H1.00.01.0.00044 </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%2Ffma84105">84105</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/Q2116409" class="extiw" title="d:Q2116409">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p><b>Pyramidal cells</b>, or <b>pyramidal neurons</b>, are a type of <a href="/wiki/Multipolar_neuron" title="Multipolar neuron">multipolar neuron</a> found in areas of the <a href="/wiki/Brain" title="Brain">brain</a> including the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a>, the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, and the <a href="/wiki/Amygdala" title="Amygdala">amygdala</a>. Pyramidal cells are the primary excitation units of the mammalian <a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">prefrontal cortex</a> and the <a href="/wiki/Corticospinal_tract" title="Corticospinal tract">corticospinal tract</a>. One of the main structural features of the pyramidal neuron is the conic shaped <a href="/wiki/Soma_(biology)" title="Soma (biology)">soma</a>, or cell body, after which the <a href="/wiki/Neuron" title="Neuron">neuron</a> is named. Other key structural features of the pyramidal cell are a single <a href="/wiki/Axon" title="Axon">axon</a>, a large <a href="/wiki/Apical_dendrite" title="Apical dendrite">apical dendrite</a>, multiple <a href="/wiki/Basal_dendrite" title="Basal dendrite">basal dendrites</a>, and the presence of <a href="/wiki/Dendritic_spines" class="mw-redirect" title="Dendritic spines">dendritic spines</a>.<sup id="cite_ref-Megias_1-0" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Pyramidal neurons are also one of two cell types where the <a href="/wiki/Pathognomonic" title="Pathognomonic">characteristic</a> <a href="/wiki/Medical_sign" class="mw-redirect" title="Medical sign">sign</a>, <a href="/wiki/Negri_bodies" class="mw-redirect" title="Negri bodies">Negri bodies</a>, are found in <a href="/wiki/Post-mortem" class="mw-redirect" title="Post-mortem">post-mortem</a> rabies infection.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Pyramidal neurons were first discovered and studied by <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a>.<sup id="cite_ref-Elston_3-0" class="reference"><a href="#cite_note-Elston-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pablo_4-0" class="reference"><a href="#cite_note-Pablo-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Since then, studies on pyramidal neurons have focused on topics ranging from <a href="/wiki/Neuroplasticity" title="Neuroplasticity">neuroplasticity</a> to <a href="/wiki/Cognition" title="Cognition">cognition</a>. </p> <style data-mw-deduplicate="TemplateStyles:r886046785">.mw-parser-output .toclimit-2 .toclevel-1 ul,.mw-parser-output .toclimit-3 .toclevel-2 ul,.mw-parser-output .toclimit-4 .toclevel-3 ul,.mw-parser-output .toclimit-5 .toclevel-4 ul,.mw-parser-output .toclimit-6 .toclevel-5 ul,.mw-parser-output .toclimit-7 .toclevel-6 ul{display:none}</style><div class="toclimit-3"><meta property="mw:PageProp/toc" /></div> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&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:Piramidal_cell.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Piramidal_cell.svg/220px-Piramidal_cell.svg.png" decoding="async" width="220" height="448" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Piramidal_cell.svg/330px-Piramidal_cell.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c1/Piramidal_cell.svg/440px-Piramidal_cell.svg.png 2x" data-file-width="420" data-file-height="855" /></a><figcaption>A reconstruction of a pyramidal cell. Soma and dendrites are labeled in red, axon arbor in blue. (1) Soma, (2) Basal dendrite, (3) Apical dendrite, (4) Axon, (5) Collateral axon.</figcaption></figure> <p>One of the main structural features of the pyramidal neuron is the conic shaped <a href="/wiki/Soma_(biology)" title="Soma (biology)">soma</a>, or cell body, after which the <a href="/wiki/Neuron" title="Neuron">neuron</a> is named. Other key structural features of the pyramidal cell are a single <a href="/wiki/Axon" title="Axon">axon</a>, a large <a href="/wiki/Apical_dendrite" title="Apical dendrite">apical dendrite</a>, multiple <a href="/wiki/Basal_dendrite" title="Basal dendrite">basal dendrites</a>, and the presence of <a href="/wiki/Dendritic_spines" class="mw-redirect" title="Dendritic spines">dendritic spines</a>.<sup id="cite_ref-Megias_1-1" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Apical_dendrite">Apical dendrite</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=2" title="Edit section: Apical dendrite"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The apical dendrite rises from the apex of the pyramidal cell's soma. The apical dendrite is a single, long, thick dendrite that branches several times as distance from the soma increases and extends towards the cortical surface.<sup id="cite_ref-Megias_1-2" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Basal_dendrite">Basal dendrite</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=3" title="Edit section: Basal dendrite"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Basal dendrites arise from the base of the soma. The basal dendritic tree consists of three to five primary dendrites. As distance increases from the soma, the basal dendrites branch profusely.<sup id="cite_ref-Megias_1-3" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Pyramidal cells are among the largest neurons in the brain. Both in humans and rodents, pyramidal cell bodies (somas) average around 20 μm in length. Pyramidal dendrites typically range in diameter from half a micrometer to several micrometers. The length of a single dendrite is usually several hundred micrometers. Due to branching, the total dendritic length of a pyramidal cell may reach several centimeters. The pyramidal cell's axon is often even longer and extensively branched, reaching many centimeters in total length. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:GFPneuron.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/GFPneuron.png/220px-GFPneuron.png" decoding="async" width="220" height="174" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/GFPneuron.png/330px-GFPneuron.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/GFPneuron.png/440px-GFPneuron.png 2x" data-file-width="459" data-file-height="364" /></a><figcaption>Pyramidal neuron visualized by <a href="/wiki/Green_fluorescent_protein" title="Green fluorescent protein">green fluorescent protein</a> (GFP)</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Hippocampal-pyramidal-cell.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Hippocampal-pyramidal-cell.png/220px-Hippocampal-pyramidal-cell.png" decoding="async" width="220" height="216" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Hippocampal-pyramidal-cell.png/330px-Hippocampal-pyramidal-cell.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Hippocampal-pyramidal-cell.png/440px-Hippocampal-pyramidal-cell.png 2x" data-file-width="450" data-file-height="442" /></a><figcaption>A <a href="/wiki/Hippocampal" class="mw-redirect" title="Hippocampal">hippocampal</a> pyramidal cell</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Dendritic_spines">Dendritic spines</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=4" title="Edit section: Dendritic spines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Dendritic_spines" class="mw-redirect" title="Dendritic spines">Dendritic spines</a> receive most of the excitatory impulses (<a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">EPSPs</a>) that enter a pyramidal cell. Dendritic spines were first noted by Ramón y Cajal in 1888 by using <a href="/wiki/Golgi%27s_method" title="Golgi's method">Golgi's method</a>. Ramón y Cajal was also the first person to propose the physiological role of increasing the receptive surface area of the neuron. The greater the pyramidal cell's surface area, the greater the neuron's ability to process and integrate large amounts of information. Dendritic spines are absent on the soma, while the number increases away from it.<sup id="cite_ref-Pablo_4-1" class="reference"><a href="#cite_note-Pablo-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The typical apical dendrite in a rat has at least 3,000 dendritic spines. The average human apical dendrite is approximately twice the length of a rat's, so the number of dendritic spines present on a human apical dendrite could be as high as 6,000.<sup id="cite_ref-LaBerge_5-0" class="reference"><a href="#cite_note-LaBerge-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Growth_and_development">Growth and development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=5" title="Edit section: Growth and development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Differentiation">Differentiation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=6" title="Edit section: Differentiation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Pyramidal specification occurs during early development of the cerebrum. <a href="/wiki/Progenitor_cells" class="mw-redirect" title="Progenitor cells">Progenitor cells</a> are committed to the neuronal lineage in the subcortical proliferative <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a> (VZ) and the <a href="/wiki/Subventricular_zone" title="Subventricular zone">subventricular zone</a> (SVZ). Immature pyramidal cells undergo migration to occupy the <a href="/wiki/Cortical_plate" class="mw-redirect" title="Cortical plate">cortical plate</a>, where they further diversify. <a href="/wiki/Endocannabinoids" class="mw-redirect" title="Endocannabinoids">Endocannabinoids</a> (eCBs) are one class of molecules that have been shown to direct pyramidal cell development and axonal pathfinding.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Transcription_factors" class="mw-redirect" title="Transcription factors">Transcription factors</a> such as Ctip2 and Sox5 have been shown to contribute to the direction in which pyramidal neurons direct their axons.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Early_postnatal_development">Early postnatal development</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=7" title="Edit section: Early postnatal development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Pyramidal cells in rats have been shown to undergo many rapid changes during early <a href="/wiki/Postnatal" class="mw-redirect" title="Postnatal">postnatal</a> life. Between postnatal days 3 and 21, pyramidal cells have been shown to double the size of the soma, increase the length of the apical dendrite fivefold, and increase basal dendrite length thirteen-fold. Other changes include the lowering of the membrane's <a href="/wiki/Resting_potential" title="Resting potential">resting potential</a>, reduction of membrane resistance, and an increase in the peak values of <a href="/wiki/Action_potential" title="Action potential">action potentials</a>.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Signaling">Signaling</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=8" title="Edit section: Signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Like dendrites in most other neurons, the dendrites are generally the input areas of the neuron, while the axon is the neuron's output. Both axons and dendrites are highly branched. The large amount of branching allows the neuron to send and receive signals to and from many different neurons. </p><p>Pyramidal neurons, like other neurons, have numerous <a href="/wiki/Voltage-gated_ion_channel" title="Voltage-gated ion channel">voltage-gated ion channels</a>. In pyramidal cells, there is an abundance of Na<sup>+</sup>, Ca<sup>2+</sup>, and K<sup>+</sup> channels in the dendrites, and some channels in the soma.<sup id="cite_ref-Spruston2008_9-0" class="reference"><a href="#cite_note-Spruston2008-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Georgiev2020_10-0" class="reference"><a href="#cite_note-Georgiev2020-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Ion channels within pyramidal cell dendrites have different properties from the same ion channel type within the pyramidal cell soma.<sup id="cite_ref-Golding2005_11-0" class="reference"><a href="#cite_note-Golding2005-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Remy2010_12-0" class="reference"><a href="#cite_note-Remy2010-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Voltage-gated Ca<sup>2+</sup> channels in pyramidal cell dendrites are activated by subthreshold <a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">EPSPs</a> and by <a href="/wiki/Neural_backpropagation" title="Neural backpropagation">back-propagating</a> action potentials. The extent of back-propagation of action potentials within pyramidal dendrites depends upon the K<sup>+</sup> channels. K<sup>+</sup> channels in pyramidal cell dendrites provide a mechanism for controlling the amplitude of action potentials.<sup id="cite_ref-Magee_13-0" class="reference"><a href="#cite_note-Magee-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>The ability of pyramidal neurons to integrate information depends on the number and distribution of the synaptic inputs they receive. A single pyramidal cell receives about 30,000 excitatory inputs and 1700 inhibitory (<a href="/wiki/IPSP" class="mw-redirect" title="IPSP">IPSPs</a>) inputs. Excitatory (EPSPs) inputs terminate exclusively on the dendritic spines, while inhibitory (IPSPs) inputs terminate on dendritic shafts, the soma, and even the axon. Pyramidal neurons can be excited by the <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a> <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>,<sup id="cite_ref-Megias_1-4" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> and inhibited by the neurotransmitter <a href="/wiki/GABA" title="GABA">GABA</a>.<sup id="cite_ref-Megias_1-5" class="reference"><a href="#cite_note-Megias-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png/220px-Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png" decoding="async" width="220" height="171" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png/330px-Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png/440px-Layer_V_Pyramidal_cell_from_mouse_visual_cortex.png 2x" data-file-width="1075" data-file-height="838" /></a><figcaption>Synaptic inputs to a Layer V Pyramidal cell in the mouse visual cortex. Each point represents one of > 11,000 post-synaptic sites on this neuron.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Firing_classifications">Firing classifications</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=9" title="Edit section: Firing classifications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Pyramidal neurons have been classified into different subclasses based upon their firing responses to 400-1000 millisecond current pulses. These classification are RSad, RSna, and IB neurons. </p> <div class="mw-heading mw-heading4"><h4 id="RSad">RSad</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=10" title="Edit section: RSad"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>RSad pyramidal neurons, or adapting regular <a href="/wiki/Biological_neuron_model" title="Biological neuron model">spiking neurons</a>, fire with individual <a href="/wiki/Action_potential" title="Action potential">action potentials</a> (APs), which are followed by a <a href="/wiki/Hyperpolarization_(biology)" title="Hyperpolarization (biology)">hyperpolarizing</a> afterpotential. The afterpotential increases in duration which creates <a href="/wiki/Action_potential" title="Action potential">spike frequency</a> <a href="/wiki/Neural_adaptation" title="Neural adaptation">adaptation</a> (SFA) in the neuron.<sup id="cite_ref-Frances_15-0" class="reference"><a href="#cite_note-Frances-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="RSna">RSna</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=11" title="Edit section: RSna"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>RSna pyramidal neurons, or non-adapting regular spiking neurons, fire a train of action potentials after a pulse. These neurons show no signs of adaptation.<sup id="cite_ref-Frances_15-1" class="reference"><a href="#cite_note-Frances-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="IB">IB</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=12" title="Edit section: IB"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>IB pyramidal neurons, or intrinsically bursting neurons, respond to <a href="/wiki/Threshold_potential" title="Threshold potential">threshold</a> pulses with a burst of two to five rapid action potentials. IB pyramidal neurons show no adaptation.<sup id="cite_ref-Frances_15-2" class="reference"><a href="#cite_note-Frances-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Molecular_classifications">Molecular classifications</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=13" title="Edit section: Molecular classifications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are several studies showing that morphological and electric pyramidal cells properties could be deduced from gene expression measured by <a href="/wiki/Single_cell_sequencing" class="mw-redirect" title="Single cell sequencing">single cell sequencing</a>.<sup id="cite_ref-Berg_151–158_16-0" class="reference"><a href="#cite_note-Berg_151–158-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Several studies are proposing that single cell classifications in mouse<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> and human<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> neurons based on gene expression could explain various neuronal properties . Neuronal types in these classifications are split into excitatory, inhibitory and hundreds of corresponding sub-types. For example, pyramidal cells of layer 2-3 in human are classified as FREM3 type<sup id="cite_ref-Berg_151–158_16-1" class="reference"><a href="#cite_note-Berg_151–158-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> and often have a high amount of Ih-current<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> generated by <a href="/wiki/HCN_channel" title="HCN channel">HCN-channel</a>. </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=Pyramidal_cell&action=edit&section=14" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Corticospinal_tract">Corticospinal tract</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=15" title="Edit section: Corticospinal tract"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Pyramidal neurons are the primary neural cell type in the <a href="/wiki/Corticospinal_tract" title="Corticospinal tract">corticospinal tract</a>. Normal motor control depends on the development of connections between the axons in the corticospinal tract and the spinal cord. Pyramidal cell axons follow cues such as growth factors to make specific connections. With proper connections, pyramidal cells take part in the circuitry responsible for vision guided motor function.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Cognition">Cognition</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=16" title="Edit section: Cognition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Pyramidal neurons in the prefrontal cortex are implicated in cognitive ability. In mammals, the complexity of pyramidal cells increases from <a href="/wiki/Posterior_(anatomy)" class="mw-redirect" title="Posterior (anatomy)">posterior</a> to <a href="/wiki/Anterior" class="mw-redirect" title="Anterior">anterior</a> brain regions. The degree of complexity of pyramidal neurons is likely linked to the cognitive capabilities of different anthropoid species. Pyramidal cells within the prefrontal cortex appear to be responsible for processing input from the primary auditory cortex, primary somatosensory cortex, and primary visual cortex, all of which process sensory modalities.<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> These cells might also play a critical role in complex object recognition within the visual processing areas of the cortex.<sup id="cite_ref-Elston_3-1" class="reference"><a href="#cite_note-Elston-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Relative to other species, the larger cell size and complexity of pyramidal neurons, along with certain patterns of cellular organization and function, correlates with the evolution of human cognition. <sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Memory_and_learning">Memory and learning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=17" title="Edit section: Memory and learning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The hippocampus's pyramidal cells are essential for certain types of memory and learning. They form synapses that aid in the integration of synaptic voltages throughout their complex dendritic trees through interactions with <a href="/wiki/Mossy_fiber_(hippocampus)" title="Mossy fiber (hippocampus)">mossy fibers</a> from <a href="/wiki/Granule_cell" title="Granule cell">granule cells</a>. Since it affects the postsynaptic voltages produced by mossy fiber activation, the placement of <a href="/wiki/Thorny_excrescence" class="mw-redirect" title="Thorny excrescence">thorny excrescences</a> on basal and apical dendrites is important for memory formation. By enabling dynamic control of the sensitivity of CA3 pyramidal cells, this clustering of mossy fiber synapses on pyramidal cells may facilitate the initiation of somatic spikes. </p><p>The interactions between pyramidal cells and an estimated 41 mossy fiber boutons, each originating from a unique granule cell, highlight the role of these boutons in information processing and synaptic connectivity, which are essential for memory and learning. Fundamentally, mossy fiber input is received by pyramidal cells in the hippocampus which integrate synaptic voltages within their dendritic architecture. The location of prickly protrusions and the clustering of synapses influence sensitivity and contribute to the processing of information pertaining to memory and learning.<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> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Pyramidal_tract" class="mw-redirect" title="Pyramidal tract">Pyramidal tract</a></li> <li><a href="/wiki/Chandelier_cell" title="Chandelier cell">Chandelier cells</a> - innervate initial segments of pyramidal axons</li> <li><a href="/wiki/Rosehip_neuron" title="Rosehip neuron">Rosehip neuron</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=19" 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 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q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Comparative+Neurology&rft.atitle=Distribution+of+thorny+excrescences+on+CA3+pyramidal+neurons+in+the+rat+hippocampus&rft.volume=430&rft.issue=3&rft.pages=357-368&rft.date=2001-02-12&rft.issn=0021-9967&rft_id=info%3Apmid%2F11169473&rft_id=info%3Adoi%2F10.1002%2F1096-9861%2820010212%29430%3A3%3C357%3A%3Aaid-cne1036%3E3.0.co%3B2-k&rft.aulast=Gonzales&rft.aufirst=R.+B.&rft.au=DeLeon+Galvan%2C+C.+J.&rft.au=Rangel%2C+Y.+M.&rft.au=Claiborne%2C+B.+J.&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F11169473%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3APyramidal+cell" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Pyramidal_cell&action=edit&section=20" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://ccdb.ucsd.edu/sand/main?mpid=51&event=displaySum">Pyramidal cell - Cell Centered Database</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20180526085757/http://static.howstuffworks.com/gif/brain-neuron-types.gif">Diagram</a></li> <li><a rel="nofollow" class="external text" href="https://wayback.archive-it.org/all/20081217005702/http://neuromedia.neurobio.ucla.edu/campbell/nervous/wp_images%5C198_pyramidal_cells.gif">Image</a></li> <li><a rel="nofollow" class="external text" href="http://www.trinity.edu/rblyston/MicroA/Lectures/L16pt1-html/sld007.htm">Diagram (as part of slideshow)</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20161102003448/http://www.trinity.edu/rblyston/MicroA/Lectures/L16pt1-html/sld007.htm">Archived</a> 2016-11-02 at the <a 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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:Nervous_tissue" title="Template:Nervous tissue"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nervous_tissue" title="Template talk:Nervous tissue"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nervous_tissue" title="Special:EditPage/Template:Nervous tissue"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Nervous_tissue" style="font-size:114%;margin:0 4em"><a href="/wiki/Nervous_tissue" title="Nervous tissue">Nervous tissue</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Central_nervous_system" title="Central nervous system">CNS</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%">Tissue Types</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/Grey_matter" title="Grey matter">Grey matter</a></li> <li><a href="/wiki/White_matter" title="White matter">White matter</a> <ul><li><a href="/wiki/Projection_fiber" title="Projection fiber">Projection fibers</a></li> <li><a href="/wiki/Association_fiber" title="Association fiber">Association fiber</a></li> <li><a href="/wiki/Commissural_fiber" title="Commissural fiber">Commissural fiber</a></li> <li><a href="/wiki/Lemniscus_(anatomy)" title="Lemniscus (anatomy)">Lemniscus</a></li> <li><a href="/wiki/Nerve_tract" title="Nerve tract">Nerve tract</a></li> <li><a href="/wiki/Decussation" title="Decussation">Decussation</a></li></ul></li> <li><a href="/wiki/Neuropil" title="Neuropil">Neuropil</a></li> <li><a href="/wiki/Meninges" title="Meninges">Meninges</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cell Types</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/Neuron" title="Neuron">Neuronal</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 class="mw-selflink selflink">Pyramidal</a></li> <li><a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje</a></li> <li><a href="/wiki/Granule_cell" title="Granule cell">Granule</a></li> <li><a href="/wiki/Von_Economo_neuron" title="Von Economo neuron">Von Economo</a></li> <li><a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">Medium spiny</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Glial</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/Astrocyte" title="Astrocyte">Astrocyte</a></li> <li><a href="/wiki/Ependyma" title="Ependyma">Ependymal cells</a> <ul><li><a href="/wiki/Tanycyte" title="Tanycyte">Tanycyte</a></li></ul></li> <li><a href="/wiki/Oligodendrocyte_progenitor_cell" title="Oligodendrocyte progenitor cell">Oligodendrocyte progenitor cell</a></li> <li><a href="/wiki/Oligodendrocyte" title="Oligodendrocyte">Oligodendrocyte</a></li> <li><a href="/wiki/Microglia" title="Microglia">Microglia</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Peripheral_nervous_system" title="Peripheral nervous system">PNS</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%">General</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i>Dorsal</i> <ul><li><a href="/wiki/Dorsal_root_of_spinal_nerve" title="Dorsal root of spinal nerve">Root</a></li> <li><a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">Ganglion</a></li> <li><a href="/wiki/Dorsal_ramus_of_spinal_nerve" title="Dorsal ramus of spinal nerve">Ramus</a></li></ul></li> <li><i>Ventral</i> <ul><li><a href="/wiki/Ventral_root_of_spinal_nerve" title="Ventral root of spinal nerve">Root</a></li> <li><a href="/wiki/Ventral_ramus_of_spinal_nerve" title="Ventral ramus of spinal nerve">Ramus</a></li></ul></li> <li><i><a href="/wiki/Ramus_communicans" title="Ramus communicans">Ramus communicans</a></i> <ul><li><a href="/wiki/Gray_ramus_communicans" title="Gray ramus communicans">Gray</a></li> <li><a href="/wiki/White_ramus_communicans" title="White ramus communicans">White</a></li></ul></li> <li><a href="/wiki/Autonomic_ganglion" title="Autonomic ganglion">Autonomic ganglion</a> (<a href="/wiki/Preganglionic_nerve_fibers" title="Preganglionic nerve fibers">Preganglionic nerve fibers</a></li> <li><a href="/wiki/Postganglionic_nerve_fibers" title="Postganglionic nerve fibers">Postganglionic nerve fibers</a>)</li> <li><a href="/wiki/Nerve_fascicle" title="Nerve fascicle">Nerve fascicle</a></li> <li><a href="/wiki/Funiculus_(neuroanatomy)" title="Funiculus (neuroanatomy)">Funiculus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissues</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/Epineurium" title="Epineurium">Epineurium</a></li> <li><a href="/wiki/Perineurium" title="Perineurium">Perineurium</a></li> <li><a href="/wiki/Endoneurium" title="Endoneurium">Endoneurium</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Neuroglia</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/Myelin" title="Myelin">Myelination</a>: <a href="/wiki/Schwann_cell" title="Schwann cell">Schwann cell</a> <ul><li><a href="/wiki/Neurilemma" title="Neurilemma">Neurilemma</a></li> <li><a href="/wiki/Myelin_incisure" title="Myelin incisure">Myelin incisure</a></li> <li><a href="/wiki/Node_of_Ranvier" title="Node of Ranvier">Node of Ranvier</a></li> <li><a href="/wiki/Internodal_segment" title="Internodal segment">Internodal segment</a></li></ul></li> <li><a href="/wiki/Satellite_glial_cell" title="Satellite glial cell">Satellite glial cell</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/Neuron" title="Neuron">Neurons</a>/<br /><a href="/wiki/Nerve_fiber" class="mw-redirect" title="Nerve fiber">nerve fibers</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%">Parts</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/Soma_(biology)" title="Soma (biology)">Soma</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/Axon_hillock" title="Axon hillock">Axon hillock</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Axon" title="Axon">Axon</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/Telodendron" class="mw-redirect" title="Telodendron">Telodendron</a></li> <li><a href="/wiki/Axon_terminal" title="Axon terminal">Axon terminals</a></li> <li><a href="/wiki/Axoplasm" title="Axoplasm">Axoplasm</a></li> <li><a href="/wiki/Axolemma" title="Axolemma">Axolemma</a></li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofibril/neurofilament</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dendrite" title="Dendrite">Dendrite</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><ul><li><a href="/wiki/Nissl_body" title="Nissl body">Nissl body</a></li> <li><a href="/wiki/Dendritic_spine" title="Dendritic spine">Dendritic spine</a></li> <li><a href="/wiki/Apical_dendrite" title="Apical dendrite">Apical dendrite</a>/<a href="/wiki/Basal_dendrite" title="Basal dendrite">Basal dendrite</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</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/Bipolar_neuron" title="Bipolar neuron">Bipolar</a></li> <li><a href="/wiki/Unipolar_neuron" title="Unipolar neuron">Unipolar</a></li> <li><a href="/wiki/Pseudounipolar_neuron" title="Pseudounipolar neuron">Pseudounipolar</a></li> <li><a href="/wiki/Multipolar_neuron" title="Multipolar neuron">Multipolar</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a> <ul><li><a href="/wiki/Renshaw_cell" title="Renshaw cell">Renshaw</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">Afferent nerve fiber</a>/<br /><a href="/wiki/Sensory_neuron" title="Sensory neuron">Sensory neuron</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/General_somatic_afferent_fibers" class="mw-redirect" title="General somatic afferent fibers">GSA</a></li> <li><a href="/wiki/General_visceral_afferent_fibers" class="mw-redirect" title="General visceral afferent fibers">GVA</a></li> <li><a href="/wiki/Special_somatic_afferent" class="mw-redirect" title="Special somatic afferent">SSA</a></li> <li><a href="/wiki/Special_visceral_afferent" class="mw-redirect" title="Special visceral afferent">SVA</a></li> <li><a href="/wiki/Axon#types" title="Axon">fibers</a> <ul><li><a href="/wiki/Type_Ia_sensory_fiber" title="Type Ia sensory fiber">Ia or Aα</a></li> <li><a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Ib or Golgi or Aα</a></li> <li><a href="/wiki/Type_II_sensory_fiber" title="Type II sensory fiber">II or Aβ and Aγ</a></li> <li><a href="/wiki/Group_A_nerve_fiber" title="Group A nerve fiber">III or Aδ or fast pain</a></li> <li><a href="/wiki/Group_C_nerve_fiber" title="Group C nerve fiber">IV or C or slow pain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">Efferent nerve fiber</a>/<br /><a href="/wiki/Motor_neuron" title="Motor neuron">Motor neuron</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/General_somatic_efferent_fibers" class="mw-redirect" title="General somatic efferent fibers">GSE</a></li> <li><a href="/wiki/General_visceral_efferent_fibers" class="mw-redirect" title="General visceral efferent fibers">GVE</a></li> <li><a href="/wiki/Special_visceral_efferent" class="mw-redirect" title="Special visceral efferent">SVE</a></li> <li><a href="/wiki/Upper_motor_neuron" title="Upper motor neuron">Upper motor neuron</a></li> <li><a href="/wiki/Lower_motor_neuron" title="Lower motor neuron">Lower motor neuron</a> <ul><li><a href="/wiki/Alpha_motor_neuron" title="Alpha motor neuron">α motorneuron</a></li> <li><a href="/wiki/Beta_motor_neuron" title="Beta motor neuron">β motorneuron</a></li> <li><a href="/wiki/Gamma_motor_neuron" title="Gamma motor neuron">γ motorneuron</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Termination</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/Synapse" title="Synapse">Synapse</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/Electrical_synapse" title="Electrical synapse">Electrical synapse</a>/<a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a> <ul><li><a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">Synaptic vesicle</a></li> <li><a href="/wiki/Active_zone" title="Active zone">Active zone</a></li> <li><a href="/wiki/Postsynaptic_density" title="Postsynaptic density">Postsynaptic density</a></li></ul></li> <li><a href="/wiki/Autapse" title="Autapse">Autapse</a></li> <li><a href="/wiki/Ribbon_synapse" title="Ribbon synapse">Ribbon synapse</a></li> <li><a href="/wiki/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sensory_receptor" class="mw-redirect" title="Sensory receptor">Sensory receptors</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/Tactile_corpuscle" title="Tactile corpuscle">Meissner's corpuscle</a></li> <li><a href="/wiki/Merkel_nerve_ending" title="Merkel nerve ending">Merkel nerve ending</a></li> <li><a href="/wiki/Pacinian_corpuscle" title="Pacinian corpuscle">Pacinian corpuscle</a></li> <li><a href="/wiki/Bulbous_corpuscle" title="Bulbous corpuscle">Ruffini ending</a></li> <li><a href="/wiki/Muscle_spindle" title="Muscle spindle">Muscle spindle</a></li> <li><a href="/wiki/Free_nerve_ending" title="Free nerve ending">Free nerve ending</a></li> <li><a href="/wiki/Nociceptor" title="Nociceptor">Nociceptor</a></li> <li><a href="/wiki/Olfactory_receptor_neuron" title="Olfactory receptor neuron">Olfactory receptor neuron</a></li> <li><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor cell</a></li> <li><a href="/wiki/Hair_cell" title="Hair cell">Hair cell</a></li> <li><a href="/wiki/Taste_receptor" title="Taste receptor">Taste receptor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved 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