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class="vector-toc-list"> <li id="toc-Glycocalyx" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Glycocalyx"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Glycocalyx</span> </div> </a> <ul id="toc-Glycocalyx-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</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-Destruction" 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Available in 26 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-26" 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">26 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%B2%D8%BA%D9%8A%D8%A8%D8%A9" title="زغيبة – Arabic" lang="ar" hreflang="ar" data-title="زغيبة" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Microvil%C2%B7li" title="Microvil·li – Catalan" lang="ca" hreflang="ca" data-title="Microvil·li" 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/Mikroklk" title="Mikroklk – Czech" lang="cs" hreflang="cs" data-title="Mikroklk" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Mikrovilli" title="Mikrovilli – German" lang="de" hreflang="de" data-title="Mikrovilli" 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/Microvilli" title="Microvilli – Spanish" lang="es" hreflang="es" data-title="Microvilli" 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%B1%DB%8C%D8%B2%D9%BE%D8%B1%D8%B2" title="ریزپرز – Persian" lang="fa" hreflang="fa" data-title="ریزپرز" 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/Microvillosit%C3%A9" title="Microvillosité – French" lang="fr" hreflang="fr" data-title="Microvillosité" 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/Microvilli" title="Microvilli – Galician" lang="gl" hreflang="gl" data-title="Microvilli" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Mikroresice" title="Mikroresice – Croatian" lang="hr" hreflang="hr" data-title="Mikroresice" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Microvilli" title="Microvilli – Italian" lang="it" hreflang="it" data-title="Microvilli" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D2%B0%D1%81%D0%B0%D2%9B%D0%B1%D2%AF%D1%80%D0%BB%D0%B5%D1%80" title="Ұсақбүрлер – Kazakh" lang="kk" hreflang="kk" data-title="Ұсақбүрлер" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Mikrogaureliai" title="Mikrogaureliai – Lithuanian" lang="lt" hreflang="lt" data-title="Mikrogaureliai" 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/Microvillus" title="Microvillus – Dutch" lang="nl" hreflang="nl" data-title="Microvillus" 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%BE%AE%E7%B5%A8%E6%AF%9B" title="微絨毛 – Japanese" lang="ja" hreflang="ja" data-title="微絨毛" 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/Mikrokosmek" title="Mikrokosmek – Polish" lang="pl" hreflang="pl" data-title="Mikrokosmek" 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/Microvilosidade" title="Microvilosidade – Portuguese" lang="pt" hreflang="pt" data-title="Microvilosidade" 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/Microvilus" title="Microvilus – Romanian" lang="ro" hreflang="ro" data-title="Microvilus" 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%9C%D0%B8%D0%BA%D1%80%D0%BE%D0%B2%D0%BE%D1%80%D1%81%D0%B8%D0%BD%D0%BA%D0%B0" title="Микроворсинка – Russian" lang="ru" hreflang="ru" data-title="Микроворсинка" 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/Microvillus" title="Microvillus – Simple English" lang="en-simple" hreflang="en-simple" data-title="Microvillus" 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-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mikrovilus" title="Mikrovilus – Slovenian" lang="sl" hreflang="sl" data-title="Mikrovilus" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Mikrovillus" title="Mikrovillus – Finnish" lang="fi" hreflang="fi" data-title="Mikrovillus" 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/Mikrovilli" title="Mikrovilli – Swedish" lang="sv" hreflang="sv" data-title="Mikrovilli" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Mikrovillus" title="Mikrovillus – Turkish" lang="tr" hreflang="tr" data-title="Mikrovillus" 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%9C%D1%96%D0%BA%D1%80%D0%BE%D0%B2%D0%BE%D1%80%D1%81%D0%B8%D0%BD%D0%BA%D0%B8" title="Мікроворсинки – Ukrainian" lang="uk" hreflang="uk" data-title="Мікроворсинки" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Vi_nhung_mao" title="Vi nhung mao – Vietnamese" lang="vi" hreflang="vi" data-title="Vi nhung mao" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li 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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">Microscopic protrusion of a cell membrane that increases surface area substantially</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:beige">Microvillus</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Human_jejunum_microvilli_2_-_TEM.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Human_jejunum_microvilli_2_-_TEM.jpg/250px-Human_jejunum_microvilli_2_-_TEM.jpg" decoding="async" width="250" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Human_jejunum_microvilli_2_-_TEM.jpg/375px-Human_jejunum_microvilli_2_-_TEM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Human_jejunum_microvilli_2_-_TEM.jpg/500px-Human_jejunum_microvilli_2_-_TEM.jpg 2x" data-file-width="640" data-file-height="480" /></a></span><div class="infobox-caption">Enterocytes with microvilli</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></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>microvillus</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=D008871">D008871</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">H1.00.01.1.01011 </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%2Ffma67296">67296</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_microanatomy" title="Anatomical terms of microanatomy">Anatomical terms of microanatomy</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q939486" class="extiw" title="d:Q939486">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-More_citations_needed plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Microvillus" title="Special:EditPage/Microvillus">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Microvillus%22">"Microvillus"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Microvillus%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Microvillus%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Microvillus%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Microvillus%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Microvillus%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">March 2024</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p><b>Microvilli</b> (<abbr title="singular form">sg.</abbr>: <b>microvillus</b>) are microscopic cellular <a href="/wiki/Membrane_protrusion" class="mw-redirect" title="Membrane protrusion">membrane protrusions</a> that increase the surface area for diffusion and minimize any increase in volume,<sup id="cite_ref-William2005_1-0" class="reference"><a href="#cite_note-William2005-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> and are involved in a wide variety of functions, including <a href="/wiki/Absorption_(chemistry)" title="Absorption (chemistry)">absorption</a>, <a href="/wiki/Secretion" title="Secretion">secretion</a>, <a href="/wiki/Cellular_adhesion" class="mw-redirect" title="Cellular adhesion">cellular adhesion</a>, and <a href="/wiki/Mechanotransduction" title="Mechanotransduction">mechanotransduction</a>. </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=Microvillus&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Microvilli are covered in plasma membrane, which encloses <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> and <a href="/wiki/Microfilaments" class="mw-redirect" title="Microfilaments">microfilaments</a>. Though these are cellular extensions, there are little or no cellular <a href="/wiki/Organelles" class="mw-redirect" title="Organelles">organelles</a> present in the microvilli. </p><p>Each microvillus has a dense bundle of cross-linked <a href="/wiki/Actin" title="Actin">actin</a> filaments, which serves as its structural core. 20 to 30 tightly bundled actin filaments are cross-linked by bundling proteins <a href="/wiki/Fimbrin" title="Fimbrin">fimbrin</a> (or plastin-1), <a href="/wiki/Villin" class="mw-redirect" title="Villin">villin</a> and <a href="/wiki/Espin_(protein)" title="Espin (protein)">espin</a> to form the core of the microvilli. </p><p>In the enterocyte microvillus, the structural core is attached to the plasma membrane along its length by lateral arms made of <a href="/wiki/MYO1A" class="mw-redirect" title="MYO1A">myosin 1a</a> and Ca<sup>2+</sup> binding protein <a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a>. Myosin 1a functions through a binding site for filamentous actin on one end and a lipid binding domain on the other. The plus ends of the actin filaments are located at the tip of the microvillus and are capped, possibly by <a href="/wiki/CapZ" title="CapZ">capZ</a> proteins,<sup id="cite_ref-books.google.co.il_2-0" class="reference"><a href="#cite_note-books.google.co.il-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> while the minus ends are anchored in the <a href="/wiki/Terminal_web" title="Terminal web">terminal web</a> composed of a complicated set of proteins including <a href="/wiki/Spectrin" title="Spectrin">spectrin</a> and myosin II. </p><p>The space between microvilli at a cell's surface is called the intermicrovillous space. Intermicrovillous space increases with contractile activity of myosin II and <a href="/wiki/Tropomyosin" title="Tropomyosin">tropomyosin</a>, and decreases when contraction ceases. </p> <div class="mw-heading mw-heading2"><h2 id="Locations">Locations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Microvillus&action=edit&section=2" title="Edit section: Locations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Thousands of microvilli form a structure called the <a href="/wiki/Brush_border" title="Brush border">brush border</a> that is found on the <a href="/wiki/Apical_membrane" class="mw-redirect" title="Apical membrane">apical surface</a> of some <a href="/wiki/Epithelial_cells" class="mw-redirect" title="Epithelial cells">epithelial cells</a>, such as the <a href="/wiki/Small_intestines" class="mw-redirect" title="Small intestines">small intestines</a>. (Microvilli should not be confused with <a href="/wiki/Intestinal_villi" class="mw-redirect" title="Intestinal villi">intestinal villi</a>, which are made of many cells. Each of these cells has many microvilli.) Microvilli are observed on the plasma surface of eggs, aiding in the anchoring of sperm cells that have penetrated the extracellular coat of egg cells. Clustering of elongated microtubules around a sperm allows for it to be drawn closer and held firmly so fusion can occur. They are large objects that increase surface area for absorption. </p><p>Microvilli are also of importance on the cell surface of <a href="/wiki/White_blood_cells" class="mw-redirect" title="White blood cells">white blood cells</a>, as they aid in the migration of white blood cells. </p> <div class="mw-heading mw-heading2"><h2 id="Relationship_to_cell">Relationship to cell</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Microvillus&action=edit&section=3" title="Edit section: Relationship to cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Microvilli are formed as cell extensions from the plasma membrane surface. </p><p>Actin filaments, present in the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>, are most abundant near the cell surface. These filaments are thought to determine the shape and movement of the plasma membrane. </p><p>The <a href="/wiki/Nucleation" title="Nucleation">nucleation</a> of actin fibers occurs as a response to external stimuli, allowing a cell to alter its shape to suit a particular situation. </p><p>This could account for the uniformity of the microvilli, which are observed to be of equal length and diameter. This nucleation process occurs from the minus end, allowing rapid growth from the plus end. </p><p>Though the length and composition of microvilli is consistent within a certain group of homogenous cells, it can differ slightly in a different part of the same organism. </p><p>For example, the microvilli in the small and large intestines in mice are slightly different in length and amount of surface coat covering.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<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=Microvillus&action=edit&section=4" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Microvilli function as the primary surface of nutrient absorption in the gastrointestinal tract. Because of this vital function, the microvillar membrane is packed with enzymes that aid in the breakdown of complex nutrients into simpler compounds that are more easily absorbed. For example, enzymes that digest carbohydrates called <a href="/wiki/Glycosidases" class="mw-redirect" title="Glycosidases">glycosidases</a> are present at high concentrations on the surface of enterocyte microvilli. Thus, microvilli not only increase the cellular surface area for absorption, they also increase the number of digestive enzymes that can be present on the cell surface. Microvilli are also present on immune cells, allowing the immune cells to sense features on the surface of pathogens and other antigen-presenting cells.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Glycocalyx">Glycocalyx</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Microvillus&action=edit&section=5" title="Edit section: Glycocalyx"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The microvilli are covered with <a href="/wiki/Glycocalyx" title="Glycocalyx">glycocalyx</a>, consisting of peripheral <a href="/wiki/Glycoproteins" class="mw-redirect" title="Glycoproteins">glycoproteins</a> that can attach themselves to a <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> via <a href="/wiki/Transmembrane_proteins" class="mw-redirect" title="Transmembrane proteins">transmembrane proteins</a>. </p><p>This layer may be used to aid binding of substances needed for uptake, to adhere nutrients or as protection against harmful elements. </p><p>It can be another location for functional enzymes to be localized. </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=Microvillus&action=edit&section=6" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Destruction">Destruction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Microvillus&action=edit&section=7" title="Edit section: Destruction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The destruction of microvilli can occur in certain diseases because of the rearrangement of cytoskeleton in host cell. This is seen in infections caused by EPEC subgroup <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i>, in <a href="/wiki/Coeliac_disease" title="Coeliac disease">celiac</a> disease, and microvillus inclusion disease<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> (an inherited disease characterized by defective microvilli and presence of cytoplasmic inclusions of the cell membrane other than the apical surface). </p><p>The destruction of microvilli can actually be beneficial sometimes, as in the case of elimination of microvilli on white blood cells which can be used to combat auto immune diseases.<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> </p><p>Congenital lack of microvilli in the intestinal tract causes <a href="/wiki/Microvillus_inclusion_disease" class="mw-redirect" title="Microvillus inclusion disease">microvillus atrophy</a>, a rare, usually fatal condition found in new-born babies. </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=Microvillus&action=edit&section=8" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Chorionic_villi" title="Chorionic villi">Chorionic villi</a></li> <li><a href="/wiki/Brush_border" title="Brush border">Brush border</a></li> <li><a href="/wiki/Cilia" class="mw-redirect" title="Cilia">Cilia</a></li> <li><a href="/wiki/Flagellum" title="Flagellum">Flagellum</a></li> <li><a href="/wiki/Intestinal_villus" title="Intestinal villus">Intestinal villus</a></li> <li><a href="/wiki/Stereocilia" title="Stereocilia">Stereocilia</a></li> <li><a href="/wiki/Terminal_web" title="Terminal web">Terminal web</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=Microvillus&action=edit&section=9" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-William2005-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-William2005_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 id="CITEREFKrause_J._William2005" class="citation book cs1">Krause J. William (July 2005). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=cRayoldYrcUC&pg=PA37"><i>Krause's Essential Human Histology for Medical Students</i></a>. Universal-Publishers. pp. 37–. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-58112-468-2" title="Special:BookSources/978-1-58112-468-2"><bdi>978-1-58112-468-2</bdi></a><span class="reference-accessdate">. Retrieved <span class="nowrap">25 November</span> 2010</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Krause%27s+Essential+Human+Histology+for+Medical+Students&rft.pages=37-&rft.pub=Universal-Publishers&rft.date=2005-07&rft.isbn=978-1-58112-468-2&rft.au=Krause+J.+William&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DcRayoldYrcUC%26pg%3DPA37&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMicrovillus" class="Z3988"></span></span> </li> <li id="cite_note-books.google.co.il-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-books.google.co.il_2-0">^</a></b></span> <span class="reference-text">The Cytoskeleton: Cytoskeleton in specialized tissues and in pathological states, edited by JE Hesketh and IF Pryme, Elsevier, 1996ISBN 1559386894, 9781559386890. Vol3, pp371, <a rel="nofollow" class="external text" href="https://books.google.com/books?id=lyrFRieEw-EC&dq=actin+filaments+capping+in+microvilli+tips&pg=PA371">M1</a><sup class="noprint Inline-Template"><span style="white-space: nowrap;">[<i><a href="/wiki/Wikipedia:Link_rot" title="Wikipedia:Link rot"><span title=" Dead link tagged November 2016">permanent dead link</span></a></i><span style="visibility:hidden; color:transparent; padding-left:2px">‍</span>]</span></sup></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMukherjee_T,_Williams_A1967" class="citation journal cs1">Mukherjee T, Williams A (1967). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2107317">"A comparative study of the ultrastructure of microvilli in the epithelium of small and large intestine of mice"</a>. <i>J Cell Biol</i>. <b>34</b> (2): 447–61. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1083%2Fjcb.34.2.447">10.1083/jcb.34.2.447</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2107317">2107317</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/6035639">6035639</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J+Cell+Biol&rft.atitle=A+comparative+study+of+the+ultrastructure+of+microvilli+in+the+epithelium+of+small+and+large+intestine+of+mice.&rft.volume=34&rft.issue=2&rft.pages=447-61&rft.date=1967&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2107317%23id-name%3DPMC&rft_id=info%3Apmid%2F6035639&rft_id=info%3Adoi%2F10.1083%2Fjcb.34.2.447&rft.au=Mukherjee+T%2C+Williams+A&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2107317&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMicrovillus" class="Z3988"></span> <a rel="nofollow" class="external text" href="http://www.jcb.org/cgi/reprint/34/2/447.pdf">link</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080406073400/http://www.jcb.org/cgi/reprint/34/2/447.pdf">Archived</a> April 6, 2008, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArameshStoychevaSanduIhle2021" class="citation journal cs1">Aramesh, Morteza; Stoycheva, Diana; Sandu, Ioana; Ihle, Stephan J.; Zünd, Tamara; Shiu, Jau-Ye; Forró, Csaba; Asghari, Mohammad; Bernero, Margherita; Lickert, Sebastian; Oxenius, Annette (2021-10-05). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8501847">"Nanoconfinement of microvilli alters gene expression and boosts T cell activation"</a>. <i>Proceedings of the National Academy of Sciences</i>. <b>118</b> (40). <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2021PNAS..11807535A">2021PNAS..11807535A</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.2107535118">10.1073/pnas.2107535118</a></span>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0027-8424">0027-8424</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8501847">8501847</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34599101">34599101</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences&rft.atitle=Nanoconfinement+of+microvilli+alters+gene+expression+and+boosts+T+cell+activation&rft.volume=118&rft.issue=40&rft.date=2021-10-05&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8501847%23id-name%3DPMC&rft_id=info%3Abibcode%2F2021PNAS..11807535A&rft_id=info%3Apmid%2F34599101&rft_id=info%3Adoi%2F10.1073%2Fpnas.2107535118&rft.issn=0027-8424&rft.aulast=Aramesh&rft.aufirst=Morteza&rft.au=Stoycheva%2C+Diana&rft.au=Sandu%2C+Ioana&rft.au=Ihle%2C+Stephan+J.&rft.au=Z%C3%BCnd%2C+Tamara&rft.au=Shiu%2C+Jau-Ye&rft.au=Forr%C3%B3%2C+Csaba&rft.au=Asghari%2C+Mohammad&rft.au=Bernero%2C+Margherita&rft.au=Lickert%2C+Sebastian&rft.au=Oxenius%2C+Annette&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8501847&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMicrovillus" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text">Malathy Kapali, MD, Ronald Jaffe, MD and Rocco M Agostini Jr. B.Sc. Final Diagnosis: Microvillus Inclusion Disease. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/19991006051512/http://path.upmc.edu/cases/case163/dx.html">"Final Diagnosis -- Case 163"</a>. Archived from <a rel="nofollow" class="external text" href="http://path.upmc.edu/cases/case163/dx.html">the original</a> on 1999-10-06<span class="reference-accessdate">. Retrieved <span class="nowrap">2006-02-19</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Final+Diagnosis+--+Case+163&rft_id=http%3A%2F%2Fpath.upmc.edu%2Fcases%2Fcase163%2Fdx.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMicrovillus" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text">Shattuck, T. (2004) Cells studied for immune function. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20160820194017/http://www.dartmouth.edu/~vox/0405/0927/cells.html">"Cells studied for immune function"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.dartmouth.edu/~vox/0405/0927/cells.html">the original</a> on 2016-08-20<span class="reference-accessdate">. Retrieved <span class="nowrap">2006-02-19</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Cells+studied+for+immune+function&rft_id=http%3A%2F%2Fwww.dartmouth.edu%2F~vox%2F0405%2F0927%2Fcells.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMicrovillus" class="Z3988"></span>.</span> </li> </ol></div></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=Microvillus&action=edit&section=10" 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="https://web.archive.org/web/20081020010317/http://trc.ucdavis.edu/mjguinan/apc100/modules/TermsCells&Tissues/structures/microvilli.html">Anatomy photo: TermsCells&Tissues/structures/microvilli</a> - Comparative Organology at University of California, Davis</li> <li><a rel="nofollow" class="external text" href="https://www.bu.edu/phpbin/medlib/histology/p/21904loa.htm">Histology image: 21904loa</a> – Histology Learning System at Boston University - "Ultrastructure of the Cell: microvilli and basal enfoldings, endocytic vesicles"</li> <li><a rel="nofollow" class="external text" href="https://www.bu.edu/phpbin/medlib/histology/p/20601loa.htm">Histology image: 20601loa</a> – Histology Learning System at Boston University - "Ultrastructure of the Cell: microvillous border and Junctional Complex, oblique section"</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 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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:Epithelial_proteins" title="Template:Epithelial proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Epithelial_proteins" title="Template talk:Epithelial proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Epithelial_proteins" title="Special:EditPage/Template:Epithelial proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Proteins_of_epithelium" style="font-size:114%;margin:0 4em">Proteins of <a href="/wiki/Epithelium" title="Epithelium">epithelium</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lateral/<a href="/wiki/Cell_junction" title="Cell junction">cell–cell</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/Cell_adhesion_molecule" title="Cell adhesion molecule">Cell adhesion molecules</a>: <a href="/wiki/Adherens_junction" title="Adherens junction">Adherens junction</a> <ul><li><a href="/wiki/Cadherin" title="Cadherin">Cadherin</a></li></ul></li> <li><a href="/wiki/Desmosome" title="Desmosome">Desmosome</a> <ul><li><a href="/wiki/Desmoglein" title="Desmoglein">Desmoglein</a></li></ul></li></ul> <ul><li><a href="/wiki/Ion_channel" title="Ion channel">Ion channels</a>: <a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a>/<a href="/wiki/Connexon" title="Connexon">Connexon</a> <ul><li><a href="/wiki/Connexin" title="Connexin">Connexin</a></li></ul></li></ul> <ul><li><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a>: <a href="/wiki/Desmosome" title="Desmosome">Desmosome</a> <ul><li><a href="/wiki/Desmoplakin" title="Desmoplakin">Desmoplakin</a></li> <li><a href="/wiki/Plakoglobin" title="Plakoglobin">Plakoglobin</a></li> <li><a href="/wiki/Tonofibril" title="Tonofibril">Tonofibril</a></li></ul></li></ul> <ul><li>other <a href="/wiki/Membrane_protein" title="Membrane protein">membrane proteins</a>: <a href="/wiki/Tight_junction" title="Tight junction">Tight junction</a> <ul><li><a href="/wiki/Claudin" title="Claudin">Claudin</a></li> <li><a href="/wiki/Occludin" title="Occludin">Occludin</a></li> <li><a href="/wiki/MARVELD2" title="MARVELD2">MARVELD2</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basal/<a href="/wiki/Cell_junction" title="Cell junction">cell–matrix</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Basal_lamina" title="Basal lamina">Basal lamina</a></li> <li><a href="/wiki/Hemidesmosome" title="Hemidesmosome">Hemidesmosome</a>/<a href="/wiki/Tonofibril" title="Tonofibril">Tonofibril</a></li> <li><a href="/wiki/Focal_adhesion" title="Focal adhesion">Focal adhesion</a></li> <li><a href="/wiki/Costamere" title="Costamere">Costamere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Apical</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/Cilium" title="Cilium">Cilia</a>/<a href="/wiki/Kinocilium" title="Kinocilium">Kinocilium</a></li> <li><a class="mw-selflink selflink">Microvilli</a>/<a href="/wiki/Stereocilia" title="Stereocilia">Stereocilia</a> (<a href="/wiki/STRC" title="STRC">STRC</a>)</li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Anatomy_of_the_gastrointestinal_tract,_excluding_the_mouth" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapsed 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:Gastrointestinal_tract" title="Template:Gastrointestinal tract"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gastrointestinal_tract" title="Template talk:Gastrointestinal tract"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gastrointestinal_tract" title="Special:EditPage/Template:Gastrointestinal tract"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Anatomy_of_the_gastrointestinal_tract,_excluding_the_mouth" style="font-size:114%;margin:0 4em">Anatomy of the <a href="/wiki/Gastrointestinal_tract" title="Gastrointestinal tract">gastrointestinal tract</a>, excluding the <a href="/wiki/Human_mouth" title="Human mouth">mouth</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Gastrointestinal_tract#Upper_gastrointestinal_tract" title="Gastrointestinal tract">Upper</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pharynx" title="Pharynx">Pharynx</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/Pharyngeal_muscles" title="Pharyngeal muscles">Muscles</a></li> <li>Spaces <ul><li><a href="/wiki/Peripharyngeal_space" title="Peripharyngeal space">peripharyngeal </a></li> <li><a href="/wiki/Retropharyngeal_space" title="Retropharyngeal space">retropharyngeal </a></li> <li><a href="/wiki/Parapharyngeal_space" title="Parapharyngeal space">parapharyngeal </a></li> <li><a href="/wiki/Retrovisceral_space" title="Retrovisceral space">retrovisceral </a></li> <li><a href="/wiki/Danger_space" title="Danger space">danger</a></li> <li><a href="/wiki/Prevertebral_space" title="Prevertebral space">prevertebral</a></li></ul></li> <li><a href="/wiki/Pterygomandibular_raphe" title="Pterygomandibular raphe">Pterygomandibular raphe</a></li> <li><a href="/wiki/Pharyngeal_raphe" title="Pharyngeal raphe">Pharyngeal raphe</a></li> <li><a href="/wiki/Buccopharyngeal_fascia" title="Buccopharyngeal fascia">Buccopharyngeal fascia</a></li> <li><a href="/wiki/Pharyngobasilar_fascia" title="Pharyngobasilar fascia">Pharyngobasilar fascia</a></li> <li><a href="/wiki/Pyriform_sinus" title="Pyriform sinus">Pyriform sinus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Esophagus" title="Esophagus">Esophagus</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>Sphincters <ul><li><a href="/wiki/Esophagus#Sphincters" title="Esophagus">upper</a></li> <li><a href="/wiki/Esophagus#Sphincters" title="Esophagus">lower</a></li></ul></li> <li><a href="/wiki/Esophageal_gland" title="Esophageal gland">glands</a></li> <li><a href="/wiki/Crop_(anatomy)" title="Crop (anatomy)">crop</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Stomach" title="Stomach">Stomach</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>Curvatures <ul><li><a href="/wiki/Curvatures_of_the_stomach#Greater_curvature" title="Curvatures of the stomach">greater</a></li> <li><a href="/wiki/Curvatures_of_the_stomach#Lesser_curvature" title="Curvatures of the stomach">lesser</a></li> <li><a href="/wiki/Angular_incisure" title="Angular incisure">Angular incisure</a></li></ul></li> <li><a href="/wiki/Stomach#Sections" title="Stomach">Cardia</a></li> <li><a href="/wiki/Stomach#Structure" title="Stomach">Body</a></li> <li><a href="/wiki/Stomach#Structure" title="Stomach">Fundus</a></li> <li><a href="/wiki/Pylorus" title="Pylorus">Pylorus</a></li> <li><a href="/wiki/Gastric_mucosa" title="Gastric mucosa">Gastric mucosa</a></li> <li><a href="/wiki/Gastric_folds" title="Gastric folds">Gastric folds</a></li> <li>Microanatomy <ul><li><a href="/wiki/Gastric_pits" title="Gastric pits">Gastric pits</a></li> <li><a href="/wiki/Gastric_glands" title="Gastric glands">Gastric glands</a></li> <li><a href="/wiki/Cardiac_glands" class="mw-redirect" title="Cardiac glands">Cardiac glands</a></li> <li><a href="/wiki/Fundic_glands" class="mw-redirect" title="Fundic glands">Fundic glands</a></li> <li><a href="/wiki/Pylorus#Histology" title="Pylorus">Pyloric glands</a></li> <li><a href="/wiki/Foveolar_cell" title="Foveolar cell">Foveolar cells</a></li> <li><a href="/wiki/Parietal_cell" title="Parietal cell">Parietal cells</a></li> <li><a href="/wiki/Gastric_chief_cell" title="Gastric chief cell">Gastric chief cells</a></li> <li><a href="/wiki/Enterochromaffin-like_cell" title="Enterochromaffin-like cell">Enterochromaffin-like cells</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/Gastrointestinal_tract#Lower_gastrointestinal_tract" title="Gastrointestinal tract">Lower</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Small_intestine" title="Small intestine">Small intestine</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Microanatomy</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/Intestinal_villus" title="Intestinal villus">Intestinal villi</a> <ul><li><a class="mw-selflink selflink">Microvilli</a></li></ul></li> <li><a href="/wiki/Intestinal_gland" title="Intestinal gland">Intestinal glands</a></li> <li><a href="/wiki/Enterocyte" title="Enterocyte">Enterocytes</a></li> <li><a href="/wiki/Enteroendocrine_cell" title="Enteroendocrine cell">Enteroendocrine cells</a></li> <li><a href="/wiki/Goblet_cell" title="Goblet cell">Goblet cells</a></li> <li><a href="/wiki/Paneth_cell" title="Paneth cell">Paneth cells</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Duodenum" title="Duodenum">Duodenum</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/Suspensory_muscle_of_duodenum" title="Suspensory muscle of duodenum">Suspensory muscle</a></li> <li><a href="/wiki/Major_duodenal_papilla" title="Major duodenal papilla">Major duodenal papilla</a></li> <li><a href="/wiki/Minor_duodenal_papilla" title="Minor duodenal papilla">Minor duodenal papilla</a></li> <li><a href="/wiki/Duodenojejunal_flexure" title="Duodenojejunal flexure">Duodenojejunal flexure</a></li> <li><a href="/wiki/Brunner%27s_glands" title="Brunner's glands">Brunner's glands</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Jejunum" title="Jejunum">Jejunum</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>No substructures</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ileum" title="Ileum">Ileum</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/Ileocecal_valve" title="Ileocecal valve">Ileocecal valve</a></li> <li><a href="/wiki/Peyer%27s_patch" title="Peyer's patch">Peyer's patches</a></li> <li><a href="/wiki/Microfold_cell" title="Microfold cell">Microfold cells</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/Large_intestine" title="Large intestine">Large intestine</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cecum" title="Cecum">Cecum</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/Appendix_(anatomy)" title="Appendix (anatomy)">Appendix</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Large_intestine#Structure" title="Large intestine">Colon</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/Ascending_colon" title="Ascending colon">Ascending colon</a></li> <li><a href="/wiki/Hepatic_flexure" class="mw-redirect" title="Hepatic flexure">Hepatic flexure</a></li> <li><a href="/wiki/Transverse_colon" title="Transverse colon">Transverse colon</a></li> <li><a href="/wiki/Splenic_flexure" class="mw-redirect" title="Splenic flexure">Splenic flexure</a></li> <li><a href="/wiki/Descending_colon" title="Descending colon">Descending colon</a></li> <li><a href="/wiki/Sigmoid_colon" title="Sigmoid colon">Sigmoid colon</a></li> <li>Continuous <ul><li><a href="/wiki/Taenia_coli" title="Taenia coli">taenia coli</a></li> <li><a href="/wiki/Haustrum_(anatomy)" title="Haustrum (anatomy)">haustra</a></li> <li><a href="/wiki/Epiploic_appendix" title="Epiploic appendix">epiploic appendix</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Rectum" title="Rectum">Rectum</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/Transverse_folds_of_rectum" title="Transverse folds of rectum">Transverse folds</a></li> <li><a href="/wiki/Rectal_ampulla" class="mw-redirect" title="Rectal ampulla">Ampulla</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Anal_canal" title="Anal canal">Anal canal</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/Human_anus" title="Human anus">Anus</a></li> <li><a href="/wiki/Anal_columns" title="Anal columns">Anal columns</a></li> <li><a href="/wiki/Anal_valves" title="Anal valves">Anal valves</a></li> <li><a href="/wiki/Anal_sinuses" title="Anal sinuses">Anal sinuses</a></li> <li><a href="/wiki/Pectinate_line" title="Pectinate line">Pectinate line</a></li> <li><a href="/wiki/Internal_anal_sphincter" title="Internal anal sphincter">Internal anal sphincter</a></li> <li><a href="/wiki/Intersphincteric_groove" class="mw-redirect" title="Intersphincteric groove">Intersphincteric groove</a></li> <li><a href="/wiki/External_anal_sphincter" title="External anal sphincter">External anal sphincter</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/Gastrointestinal_wall" title="Gastrointestinal wall">Wall</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/Serous_membrane" title="Serous membrane">Serosa</a> / <a href="/wiki/Adventitia" title="Adventitia">Adventitia</a></li> <li><a href="/wiki/Subserosa" title="Subserosa">Subserosa</a></li> <li><a href="/wiki/Muscular_layer" title="Muscular layer">Muscular layer</a></li> <li><a href="/wiki/Submucosa" title="Submucosa">Submucosa</a></li> <li><a href="/wiki/Circular_folds" title="Circular folds">Circular folds</a></li> <li><a href="/wiki/Mucous_membrane" title="Mucous membrane">Mucosa</a></li> <li><a href="/wiki/Muscularis_mucosae" title="Muscularis mucosae">Muscularis mucosa</a></li></ul> </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> 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