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class="vector-toc-list"> <li id="toc-Basal_body" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Basal_body"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Basal body</span> </div> </a> <ul id="toc-Basal_body-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ciliary_rootlet" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ciliary_rootlet"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Ciliary rootlet</span> </div> </a> <ul id="toc-Ciliary_rootlet-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transition_zone" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transition_zone"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Transition zone</span> </div> </a> <ul id="toc-Transition_zone-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Axoneme" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Axoneme"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Axoneme</span> </div> </a> <ul id="toc-Axoneme-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Types"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Types</span> </div> </a> <button aria-controls="toc-Types-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 Types subsection</span> </button> <ul id="toc-Types-sublist" class="vector-toc-list"> <li id="toc-Non-motile_cilia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-motile_cilia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Non-motile cilia</span> </div> </a> <ul id="toc-Non-motile_cilia-sublist" class="vector-toc-list"> <li id="toc-Modified_non-motile_cilia" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Modified_non-motile_cilia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>Modified non-motile cilia</span> </div> </a> <ul id="toc-Modified_non-motile_cilia-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Motile_cilia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Motile_cilia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Motile cilia</span> </div> </a> <ul id="toc-Motile_cilia-sublist" class="vector-toc-list"> <li id="toc-Modified_motile_cilia" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Modified_motile_cilia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.1</span> <span>Modified motile cilia</span> </div> </a> <ul id="toc-Modified_motile_cilia-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nodal_cilia" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Nodal_cilia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.2</span> <span>Nodal cilia</span> </div> </a> <ul id="toc-Nodal_cilia-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Cilia_versus_flagella" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cilia_versus_flagella"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Cilia versus flagella</span> </div> </a> <ul id="toc-Cilia_versus_flagella-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Microorganisms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Microorganisms"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Microorganisms</span> </div> </a> <ul id="toc-Microorganisms-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Ciliogenesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Ciliogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Ciliogenesis</span> </div> </a> <ul id="toc-Ciliogenesis-sublist" class="vector-toc-list"> </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-Sensing_the_extracellular_environment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sensing_the_extracellular_environment"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Sensing the extracellular environment</span> </div> </a> <ul id="toc-Sensing_the_extracellular_environment-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Axo-ciliary_synapse" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Axo-ciliary_synapse"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Axo-ciliary synapse</span> </div> </a> <ul id="toc-Axo-ciliary_synapse-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-Extracellular_changes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Extracellular_changes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Extracellular changes</span> </div> </a> <ul id="toc-Extracellular_changes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Primary_cilia_in_pancreatic_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Primary_cilia_in_pancreatic_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Primary cilia in pancreatic cells</span> </div> </a> <ul id="toc-Primary_cilia_in_pancreatic_cells-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">6</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">7</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">8</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">Cilium</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 47 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-47" 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">47 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%87%D8%AF%D8%A8" title="هدب – Arabic" lang="ar" hreflang="ar" data-title="هدب" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A0%D0%B5%D1%81%D0%BD%D0%B8%D1%87%D0%BA%D0%B8" title="Реснички – Bulgarian" lang="bg" hreflang="bg" data-title="Реснички" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Treplja" title="Treplja – Bosnian" lang="bs" hreflang="bs" data-title="Treplja" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Cili" title="Cili – Catalan" lang="ca" hreflang="ca" data-title="Cili" 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/%C5%98asinka" title="Řasinka – Czech" lang="cs" hreflang="cs" data-title="Řasinka" 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/Fimreh%C3%A5r" title="Fimrehår – Danish" lang="da" hreflang="da" data-title="Fimrehår" 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/Zilie" title="Zilie – German" lang="de" hreflang="de" data-title="Zilie" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Ripsmed_(organellid)" title="Ripsmed (organellid) – Estonian" lang="et" hreflang="et" data-title="Ripsmed (organellid)" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Cilio" title="Cilio – Spanish" lang="es" hreflang="es" data-title="Cilio" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Cilio" title="Cilio – Esperanto" lang="eo" hreflang="eo" data-title="Cilio" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Zilio" title="Zilio – Basque" lang="eu" hreflang="eu" data-title="Zilio" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%85%DA%98%DA%A9" title="مژک – Persian" lang="fa" hreflang="fa" data-title="مژک" 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/Cil_cellulaire" title="Cil cellulaire – French" lang="fr" hreflang="fr" data-title="Cil cellulaire" 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/Cilio" title="Cilio – Galician" lang="gl" hreflang="gl" data-title="Cilio" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Cilio_(vejetanto)" title="Cilio (vejetanto) – Ido" lang="io" hreflang="io" data-title="Cilio (vejetanto)" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Silia" title="Silia – Indonesian" lang="id" hreflang="id" data-title="Silia" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Bifh%C3%A1r" title="Bifhár – Icelandic" lang="is" hreflang="is" data-title="Bifhár" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Ciglia_(biologia)" title="Ciglia (biologia) – Italian" lang="it" hreflang="it" data-title="Ciglia (biologia)" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A8%D7%99%D7%A1%D7%95%D7%9F_(%D7%91%D7%99%D7%95%D7%9C%D7%95%D7%92%D7%99%D7%94)" title="ריסון (ביולוגיה) – Hebrew" lang="he" hreflang="he" data-title="ריסון (ביולוגיה)" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Silia" title="Silia – Javanese" lang="jv" hreflang="jv" data-title="Silia" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9A%D1%96%D1%80%D0%BF%D1%96%D0%BA%D1%88%D0%B5%D0%BB%D0%B5%D1%80" title="Кірпікшелер – Kazakh" lang="kk" hreflang="kk" data-title="Кірпікшелер" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Sil" title="Sil – Haitian Creole" lang="ht" hreflang="ht" data-title="Sil" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Skropsti%C5%86as" title="Skropstiņas – Latvian" lang="lv" hreflang="lv" data-title="Skropstiņas" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Blakstien%C4%97l%C4%97" title="Blakstienėlė – Lithuanian" lang="lt" hreflang="lt" data-title="Blakstienėlė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Csill%C3%B3" title="Csilló – Hungarian" lang="hu" hreflang="hu" data-title="Csilló" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Silium" title="Silium – Malay" lang="ms" hreflang="ms" data-title="Silium" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Trilhaar" title="Trilhaar – Dutch" lang="nl" hreflang="nl" data-title="Trilhaar" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B9%8A%E6%AF%9B" title="繊毛 – Japanese" lang="ja" hreflang="ja" data-title="繊毛" 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-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Flimmerh%C3%A5r" title="Flimmerhår – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Flimmerhår" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Flimmerh%C3%A5r" title="Flimmerhår – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Flimmerhår" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Rz%C4%99ska" title="Rzęska – Polish" lang="pl" hreflang="pl" data-title="Rzęska" 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%ADlio" title="Cílio – Portuguese" lang="pt" hreflang="pt" data-title="Cílio" 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/Cil" title="Cil – Romanian" lang="ro" hreflang="ro" data-title="Cil" 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%A0%D0%B5%D1%81%D0%BD%D0%B8%D1%87%D0%BA%D0%B8" title="Реснички – Russian" lang="ru" hreflang="ru" data-title="Реснички" 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/Cilia" title="Cilia – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cilia" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/C%C3%ADlium_(bunka)" title="Cílium (bunka) – Slovak" lang="sk" hreflang="sk" data-title="Cílium (bunka)" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Migetalka" title="Migetalka – Slovenian" lang="sl" hreflang="sl" data-title="Migetalka" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A2%D1%80%D0%B5%D0%BF%D1%99%D0%B5" title="Трепље – Serbian" lang="sr" hreflang="sr" data-title="Трепље" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Treplje" title="Treplje – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Treplje" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Silia" title="Silia – Sundanese" lang="su" hreflang="su" data-title="Silia" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/V%C3%A4rekarva" title="Värekarva – Finnish" lang="fi" hreflang="fi" data-title="Värekarva" data-language-autonym="Suomi" 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class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Organelle found on eukaryotic cells</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about organelles. For fine hairs on insect wings, see <a href="/wiki/Cilium_(entomology)" class="mw-redirect" title="Cilium (entomology)">Cilium (entomology)</a>.</div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Cilium_(computing)" title="Cilium (computing)">Cilium (computing)</a>.</div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Psyllium" title="Psyllium">Psyllium</a>.</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:beige">Cilium</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Bronchiolar_epithelium_3_-_SEM.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Bronchiolar_epithelium_3_-_SEM.jpg/250px-Bronchiolar_epithelium_3_-_SEM.jpg" decoding="async" width="250" height="256" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Bronchiolar_epithelium_3_-_SEM.jpg/375px-Bronchiolar_epithelium_3_-_SEM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/39/Bronchiolar_epithelium_3_-_SEM.jpg/500px-Bronchiolar_epithelium_3_-_SEM.jpg 2x" data-file-width="1024" data-file-height="1049" /></a></span><div class="infobox-caption"><a href="/wiki/Scanning_electron_microscope" title="Scanning electron microscope">SEM</a> micrograph of motile cilia projecting from <a href="/wiki/Respiratory_epithelium" title="Respiratory epithelium">respiratory epithelium</a> in the <a href="/wiki/Trachea" title="Trachea">trachea</a></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>cilium</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=D002923">D002923</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">H1.00.01.1.01014 </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%2Ffma67181">67181</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/Q180436" class="extiw" title="d:Q180436">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/40px-Wiktionary-logo-en-v2.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/60px-Wiktionary-logo-en-v2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></span></span></div> <div class="side-box-text plainlist">Look up <i><b><a href="https://en.wiktionary.org/wiki/cilium" class="extiw" title="wiktionary:cilium"> cilium</a></b></i> in Wiktionary, the free dictionary.</div></div> </div> <p>The <b>cilium</b> (<abbr title="plural form">pl.</abbr>: <b>cilia</b>; from <a href="/wiki/Latin_language" class="mw-redirect" title="Latin language">Latin</a> <i> cilium</i> '<a href="/wiki/Eyelid" title="Eyelid">eyelid</a>'; in Medieval Latin and in anatomy, <i>cilium</i>) is a short hair-like <a href="/wiki/Membrane_protrusion" class="mw-redirect" title="Membrane protrusion">membrane protrusion</a> from many types of <a href="/wiki/Eukaryotic_cell" class="mw-redirect" title="Eukaryotic cell">eukaryotic cell</a>.<sup id="cite_ref-MW1_1-0" class="reference"><a href="#cite_note-MW1-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-HHMIB2005_2-0" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> (Cilia are absent in <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>.) The cilium has the shape of a slender threadlike projection that extends from the surface of the much larger cell body.<sup id="cite_ref-HHMIB2005_2-1" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Eukaryotic <a href="/wiki/Flagella" class="mw-redirect" title="Flagella">flagella</a> found on <a href="/wiki/Sperm_cell" class="mw-redirect" title="Sperm cell">sperm cells</a> and many <a href="/wiki/Protozoan" class="mw-redirect" title="Protozoan">protozoans</a> have a similar structure to motile cilia that enables swimming through liquids; they are longer than cilia and have a different undulating motion.<sup id="cite_ref-Haimo_JCB198112_3-0" class="reference"><a href="#cite_note-Haimo_JCB198112-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Alberts1_4-0" class="reference"><a href="#cite_note-Alberts1-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>There are two major classes of cilia: <i>motile</i> and <i>non-motile</i> cilia, each with two subtypes, giving four types in all.<sup id="cite_ref-Falk_5-0" class="reference"><a href="#cite_note-Falk-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> A cell will typically have one primary cilium or many motile cilia.<sup id="cite_ref-Wheatley_6-0" class="reference"><a href="#cite_note-Wheatley-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The structure of the cilium core, called the <a href="/wiki/Axoneme" title="Axoneme">axoneme</a>, determines the cilium class. Most motile cilia have a central pair of single <a href="/wiki/Microtubule" title="Microtubule">microtubules</a> surrounded by nine pairs of double microtubules called a <a href="/wiki/9%2B2_axoneme" class="mw-redirect" title="9+2 axoneme">9+2 axoneme</a>. Most non-motile cilia have a <a href="/wiki/9%2B0_axoneme" class="mw-redirect" title="9+0 axoneme">9+0 axoneme</a> that lacks the central pair of microtubules. Also lacking are the associated components that enable motility including the outer and inner <a href="/wiki/Dynein" title="Dynein">dynein</a> arms, and radial spokes.<sup id="cite_ref-Fisch_7-0" class="reference"><a href="#cite_note-Fisch-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Some motile cilia lack the central pair, and some non-motile cilia have the central pair, hence the four types.<sup id="cite_ref-Falk_5-1" class="reference"><a href="#cite_note-Falk-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fisch_7-1" class="reference"><a href="#cite_note-Fisch-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Most non-motile cilia, termed <i>primary cilia</i> or <i>sensory cilia</i>, serve solely as sensory organelles.<sup id="cite_ref-Prevo_8-0" class="reference"><a href="#cite_note-Prevo-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Elliott_9-0" class="reference"><a href="#cite_note-Elliott-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Most vertebrate cell types possess a single non-motile primary cilium, which functions as a cellular antenna.<sup id="cite_ref-Singla_629–633_10-0" class="reference"><a href="#cite_note-Singla_629–633-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Patel_11-0" class="reference"><a href="#cite_note-Patel-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Olfactory_neuron" class="mw-redirect" title="Olfactory neuron">Olfactory neurons</a> possess a great many non-motile cilia. Non-motile cilia that have a central pair of microtubules are the <a href="/wiki/Kinocilium" title="Kinocilium">kinocilia</a> present on <a href="/wiki/Hair_cell" title="Hair cell">hair cells</a>.<sup id="cite_ref-Falk_5-2" class="reference"><a href="#cite_note-Falk-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Motile cilia are found in large numbers on <a href="/wiki/Respiratory_epithelium" title="Respiratory epithelium">respiratory epithelial cells</a> – around 200 cilia per cell, where they function in <a href="/wiki/Mucociliary_clearance" title="Mucociliary clearance">mucociliary clearance</a>, and also have <a href="/wiki/Mechanosensation" title="Mechanosensation">mechanosensory</a> and <a href="/wiki/Chemoreceptor" title="Chemoreceptor">chemosensory</a> functions.<sup id="cite_ref-Horani_12-0" class="reference"><a href="#cite_note-Horani-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2012-Enuka_13-0" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bloodgood_14-0" class="reference"><a href="#cite_note-Bloodgood-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Motile cilia on <a href="/wiki/Ependymal_cells" class="mw-redirect" title="Ependymal cells">ependymal cells</a> move the <a href="/wiki/Cerebrospinal_fluid" title="Cerebrospinal fluid">cerebrospinal fluid</a> through the <a href="/wiki/Ventricular_system" title="Ventricular system">ventricular system</a> of the <a href="/wiki/Brain" title="Brain">brain</a>. Motile cilia are also present in the <a href="/wiki/Oviduct" title="Oviduct">oviducts</a> (<a href="/wiki/Fallopian_tube" title="Fallopian tube">fallopian tubes</a>) of female (<a href="/wiki/Theria" title="Theria">therian</a>) mammals, where they function in moving <a href="/wiki/Egg_cell" title="Egg cell">egg cells</a> from the <a href="/wiki/Ovary" title="Ovary">ovary</a> to the <a href="/wiki/Uterus" title="Uterus">uterus</a>.<sup id="cite_ref-2012-Enuka_13-1" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Panelli_15-0" class="reference"><a href="#cite_note-Panelli-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Motile cilia that lack the central pair of microtubules are found in the cells of the embryonic <a href="/wiki/Primitive_node" title="Primitive node">primitive node</a>; termed <i>nodal cells</i>, these nodal cilia are responsible for the <a href="/wiki/Left-right_asymmetry" title="Left-right asymmetry">left-right asymmetry</a> of <a href="/wiki/Bilaterians" class="mw-redirect" title="Bilaterians">bilaterians</a>.<sup id="cite_ref-Desgrange_16-0" class="reference"><a href="#cite_note-Desgrange-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Eukaryotic_cilium_diagram_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Eukaryotic_cilium_diagram_en.svg/300px-Eukaryotic_cilium_diagram_en.svg.png" decoding="async" width="300" height="256" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Eukaryotic_cilium_diagram_en.svg/450px-Eukaryotic_cilium_diagram_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Eukaryotic_cilium_diagram_en.svg/600px-Eukaryotic_cilium_diagram_en.svg.png 2x" data-file-width="505" data-file-height="431" /></a><figcaption>Eukaryotic motile cilium</figcaption></figure> <p>A cilium is assembled and built from a <a href="/wiki/Basal_body" title="Basal body">basal body</a> on the cell surface. From the basal body, the ciliary rootlet forms ahead of the transition plate and transition zone where the earlier microtubule triplets change to the microtubule doublets of the axoneme. </p> <div class="mw-heading mw-heading3"><h3 id="Basal_body">Basal body</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=2" title="Edit section: Basal body"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The foundation of the cilium is the basal body, a term applied to the mother centriole when it is associated with a cilium. Mammalian basal bodies consist of a barrel of nine triplet microtubules, subdistal appendages and nine strut-like structures, known as distal appendages, which attach the basal body to the membrane at the base of the cilium. Two of each of the basal body's triplet microtubules extend during growth of the axoneme to become the doublet microtubules. </p> <div class="mw-heading mw-heading3"><h3 id="Ciliary_rootlet">Ciliary rootlet</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=3" title="Edit section: Ciliary rootlet"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ciliary rootlet is a cytoskeleton-like structure that originates from the basal body at the proximal end of a cilium. Rootlets are typically 80-100 nm in diameter and contain cross striae distributed at regular intervals of approximately 55-70 nm. A prominent component of the rootlet is <a href="/wiki/Rootletin" title="Rootletin">rootletin</a> a coiled coil rootlet protein coded for by the <a href="/wiki/Rootletin" title="Rootletin"><i>CROCC</i> gene</a>.<sup id="cite_ref-uniprot_17-0" class="reference"><a href="#cite_note-uniprot-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transition_zone">Transition zone</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=4" title="Edit section: Transition zone"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>To achieve its distinct composition, the proximal-most region of the cilium consists of a <b>transition zone</b>, also known as the <b>ciliary gate</b>, that controls the entry and exit of proteins to and from the cilium.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><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> At the transition zone, Y-shaped structures connect the ciliary membrane to the underlying axoneme. Control of selective entry into cilia may involve a sieve-like function of transition zone. Inherited defects in components of the transition zone cause ciliopathies, such as Joubert syndrome. Transition zone structure and function is conserved across diverse organisms, including vertebrates, <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">Caenorhabditis elegans</a></i>, <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> and <i><a href="/wiki/Chlamydomonas_reinhardtii" title="Chlamydomonas reinhardtii">Chlamydomonas reinhardtii</a></i>. In mammals, disruption of the transition zone reduces the ciliary abundance of membrane-associated ciliary proteins, such as those involved in <a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signal transduction</a>, compromising Hedgehog-dependent embryonic development of digit number and central nervous system patterning. </p> <div class="mw-heading mw-heading3"><h3 id="Axoneme">Axoneme</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=5" title="Edit section: Axoneme"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Inside a cilium is a <a href="/wiki/Microtubule" title="Microtubule">microtubule</a>-based <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeletal core</a> called the <a href="/wiki/Axoneme" title="Axoneme">axoneme</a>. The axoneme of a primary cilium typically has a ring of nine outer microtubule doublets (called a <a href="/wiki/9%2B0_axoneme" class="mw-redirect" title="9+0 axoneme">9+0 axoneme</a>), and the axoneme of a motile cilium has, in addition to the nine outer doublets, two central microtubule singlets (called a <a href="/wiki/9%2B2_axoneme" class="mw-redirect" title="9+2 axoneme">9+2 axoneme</a>). This is the same axoneme type of the <a href="/wiki/Flagellum" title="Flagellum">flagellum</a>. The axoneme in a motile cilium acts as a scaffold for the inner and outer <a href="/wiki/Dynein_arm" class="mw-redirect" title="Dynein arm">dynein arms</a> that move the cilium, and provides tracks for the <a href="/wiki/Microtubule_motor" class="mw-redirect" title="Microtubule motor">microtubule motor</a> <a href="/wiki/Motor_protein" title="Motor protein">proteins</a> of kinesin and dynein.<sup id="cite_ref-HHMIB2005_2-2" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The transport of ciliary components is carried out by <a href="/wiki/Intraflagellar_transport" title="Intraflagellar transport">intraflagellar transport</a> (IFT) which is similar to the <a href="/wiki/Axonal_transport" title="Axonal transport">axonal transport</a> in a <a href="/wiki/Axon" title="Axon">nerve fibre</a>. Transport is bidirectional and <a href="/wiki/Motor_protein#Cytoskeletal_motor_proteins" title="Motor protein">cytoskeletal motor proteins</a> kinesin and dynein transport ciliary components along the microtubule tracks; kinesin in an anterograde movement towards the ciliary tip and dynein in a retrograde movement towards the cell body. The cilium has its own ciliary membrane enclosed within the surrounding <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a>.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=6" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Non-motile_cilia">Non-motile cilia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=7" title="Edit section: Non-motile cilia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In animals, non-motile <b>primary cilia</b> are found on nearly every type of cell, blood cells being a prominent exception.<sup id="cite_ref-HHMIB2005_2-3" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Most cells only possess one, in contrast to cells with motile cilia, an exception being <a href="/wiki/Olfactory_epithelium#Olfactory_sensory_neurons" title="Olfactory epithelium">olfactory sensory neurons</a>, where the <a href="/wiki/Odorant_receptors" class="mw-redirect" title="Odorant receptors">odorant receptors</a> are located, which each possess about ten cilia. Some cell types, such as retinal photoreceptor cells, possess highly specialized primary cilia.<sup id="cite_ref-Wolfrum_24-0" class="reference"><a href="#cite_note-Wolfrum-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Although the primary cilium was discovered in 1898, it was largely ignored for a century and considered a <a href="/wiki/Vestigial" class="mw-redirect" title="Vestigial">vestigial</a> organelle without important function.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-HHMIB2005_2-4" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Recent findings regarding its physiological roles in chemosensation, signal transduction, and cell growth control, have revealed its importance in cell function. Its importance to human biology has been underscored by the discovery of its role in a diverse group of diseases caused by the <a href="/wiki/Anterior_segment_dysgenesis" class="mw-redirect" title="Anterior segment dysgenesis">dysgenesis</a> or dysfunction of cilia, such as <a href="/wiki/Polycystic_kidney_disease" title="Polycystic kidney disease">polycystic kidney disease</a>,<sup id="cite_ref-pmid18264930_26-0" class="reference"><a href="#cite_note-pmid18264930-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Congenital_heart_disease" class="mw-redirect" title="Congenital heart disease">congenital heart disease</a>,<sup id="cite_ref-pmid17548739_27-0" class="reference"><a href="#cite_note-pmid17548739-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Mitral_valve_prolapse" title="Mitral valve prolapse">mitral valve prolapse</a>,<sup id="cite_ref-pmid_32380895_28-0" class="reference"><a href="#cite_note-pmid_32380895-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> and retinal degeneration,<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> called <a href="/wiki/Ciliopathy" title="Ciliopathy">ciliopathies</a>.<sup id="cite_ref-badano2006_30-0" class="reference"><a href="#cite_note-badano2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Reiter_533–547_31-0" class="reference"><a href="#cite_note-Reiter_533–547-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> The primary cilium is now known to play an important role in the function of many human organs.<sup id="cite_ref-HHMIB2005_2-5" class="reference"><a href="#cite_note-HHMIB2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Singla_629–633_10-1" class="reference"><a href="#cite_note-Singla_629–633-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Primary cilia on pancreatic <a href="/wiki/Beta_cell" title="Beta cell">beta cells</a> regulate their function and energy metabolism. Cilia deletion can lead to islet dysfunction and <a href="/wiki/Type_2_diabetes" title="Type 2 diabetes">type 2 diabetes</a>.<sup id="cite_ref-Hegyi_32-0" class="reference"><a href="#cite_note-Hegyi-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Cilia are assembled during the <a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub> phase</a> and are disassembled before mitosis occurs.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Patel_11-1" class="reference"><a href="#cite_note-Patel-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Disassembly of cilia requires the action of <a href="/wiki/Aurora_kinase_A" title="Aurora kinase A">aurora kinase A</a>.<sup id="cite_ref-entrez_34-0" class="reference"><a href="#cite_note-entrez-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> The current scientific understanding of primary cilia views them as "sensory <a href="/wiki/Cell_(biology)" title="Cell (biology)">cellular</a> antennae that coordinate many cellular signaling pathways, sometimes coupling the signaling to ciliary motility or alternatively to cell division and differentiation."<sup id="cite_ref-Satir2008_35-0" class="reference"><a href="#cite_note-Satir2008-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> The cilium is composed of subdomains<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (October 2020)">clarification needed</span></a></i>]</sup> and enclosed by a plasma membrane continuous with the plasma membrane of the cell. For many cilia, the <a href="/wiki/Basal_body" title="Basal body">basal body</a>, where the cilium originates, is located within a membrane invagination called the ciliary pocket. The cilium membrane and the basal body microtubules are connected by distal appendages (also called transition fibers). Vesicles carrying molecules for the cilia dock at the distal appendages. Distal to the transition fibers form a transition zone where entry and exit of molecules is regulated to and from the cilia. Some of the signaling with these cilia occur through ligand binding such as <a href="/wiki/Hedgehog_signaling" class="mw-redirect" title="Hedgehog signaling">Hedgehog signaling</a>.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Other forms of signaling include <a href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">G protein-coupled receptors</a> including the <a href="/wiki/Somatostatin_receptor_3" title="Somatostatin receptor 3">somatostatin receptor 3</a> in neurons.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Blausen_0766_RespiratoryEpithelium.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Blausen_0766_RespiratoryEpithelium.png/220px-Blausen_0766_RespiratoryEpithelium.png" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Blausen_0766_RespiratoryEpithelium.png/330px-Blausen_0766_RespiratoryEpithelium.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Blausen_0766_RespiratoryEpithelium.png/440px-Blausen_0766_RespiratoryEpithelium.png 2x" data-file-width="2500" data-file-height="2500" /></a><figcaption>Illustration depicting motile cilia on <a href="/wiki/Respiratory_epithelium" title="Respiratory epithelium">respiratory epithelium</a>.</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Modified_non-motile_cilia">Modified non-motile cilia</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=8" title="Edit section: Modified non-motile cilia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Kinocilia" class="mw-redirect" title="Kinocilia">Kinocilia</a> that are found on hair cells in the inner ear are termed as specialized primary cilia, or modified non-motile cilia. They possess the 9+2 axoneme of the motile cilia but lack the inner dynein arms that give movement. They do move passively following the detection of sound, allowed by the outer dynein arms.<sup id="cite_ref-Wang_38-0" class="reference"><a href="#cite_note-Wang-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Takeda_39-0" class="reference"><a href="#cite_note-Takeda-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Motile_cilia">Motile cilia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=9" title="Edit section: Motile cilia"><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:Bronchiolar_epithelium_4_-_SEM.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Bronchiolar_epithelium_4_-_SEM.jpg/220px-Bronchiolar_epithelium_4_-_SEM.jpg" decoding="async" width="220" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Bronchiolar_epithelium_4_-_SEM.jpg/330px-Bronchiolar_epithelium_4_-_SEM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Bronchiolar_epithelium_4_-_SEM.jpg/440px-Bronchiolar_epithelium_4_-_SEM.jpg 2x" data-file-width="1024" data-file-height="1049" /></a><figcaption><a href="/wiki/Trachea" title="Trachea">Tracheal</a> respiratory epithelium showing cilia and much smaller <a href="/wiki/Microvilli" class="mw-redirect" title="Microvilli">microvilli</a> on non-ciliated cells in <a href="/wiki/Scanning_electron_microscope" title="Scanning electron microscope">scanning electron micrograph</a>.</figcaption></figure> <p><a href="/wiki/Mammal" title="Mammal">Mammals</a> also have <b>motile cilia</b> or <b>secondary cilia</b> that are usually present on a cell's surface in large numbers (multiciliate), and beat in coordinated <a href="/wiki/Metachronal_rhythm" title="Metachronal rhythm">metachronal waves</a>.<sup id="cite_ref-Lewin2007_40-0" class="reference"><a href="#cite_note-Lewin2007-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Multiciliated cells are found <a href="/wiki/Respiratory_epithelium" title="Respiratory epithelium">lining the respiratory tract</a> where they function in <a href="/wiki/Mucociliary_clearance" title="Mucociliary clearance">mucociliary clearance</a> sweeping mucus containing debris away from the <a href="/wiki/Lung" title="Lung">lungs</a>.<sup id="cite_ref-2012-Enuka_13-2" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Each cell in the respiratory epithelium has around 200 motile cilia.<sup id="cite_ref-Horani_12-1" class="reference"><a href="#cite_note-Horani-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>In the <a href="/wiki/Reproductive_tract" class="mw-redirect" title="Reproductive tract">reproductive tract</a>, <a href="/wiki/Smooth_muscle" title="Smooth muscle">smooth muscle</a> contractions help the beating of the cilia in moving the <a href="/wiki/Egg_cell" title="Egg cell">egg cell</a> from the ovary to the uterus.<sup id="cite_ref-2012-Enuka_13-3" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Panelli_15-1" class="reference"><a href="#cite_note-Panelli-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/Ventricular_system" title="Ventricular system">ventricles of the brain</a> ciliated <a href="/wiki/Ependymal_cells" class="mw-redirect" title="Ependymal cells">ependymal cells</a> circulate the <a href="/wiki/Cerebrospinal_fluid" title="Cerebrospinal fluid">cerebrospinal fluid</a>. </p><p>The functioning of motile cilia is strongly dependent on the maintenance of optimal levels of <a href="/wiki/Periciliary_liquid_layer" class="mw-redirect" title="Periciliary liquid layer">periciliary fluid</a> bathing the cilia. <a href="/wiki/Epithelial_sodium_channel" title="Epithelial sodium channel">Epithelial sodium channels</a> (ENaCs) are specifically expressed along the entire length of cilia in the respiratory tract, and <a href="/wiki/Fallopian_tube" title="Fallopian tube">fallopian tube</a> or <i><a href="/wiki/Oviduct" title="Oviduct">oviduct</a></i> that apparently serve as sensors to regulate the periciliary fluid.<sup id="cite_ref-2012-Enuka_13-4" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2016-Hanukoglu_41-0" class="reference"><a href="#cite_note-2016-Hanukoglu-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Modified_motile_cilia">Modified motile cilia</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=10" title="Edit section: Modified motile cilia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Motile cilia without the central pair of singlets (9+0) are found in early embryonic development. They are present as nodal cilia on the nodal cells of the <a href="/wiki/Primitive_node" title="Primitive node">primitive node</a>. Nodal cells are responsible for the <a href="/wiki/Left-right_asymmetry" title="Left-right asymmetry">left-right asymmetry</a> in <a href="/wiki/Bilaterians" class="mw-redirect" title="Bilaterians">bilateral animals</a>.<sup id="cite_ref-Desgrange_16-1" class="reference"><a href="#cite_note-Desgrange-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> While lacking the central apparatus there are <a href="/wiki/Dynein_arm" class="mw-redirect" title="Dynein arm">dynein arms</a> present that allow the nodal cilia to move in a spinning fashion. The movement creates a current flow of the extraembryonic fluid across the <a href="/wiki/Primitive_node" title="Primitive node">nodal surface</a> in a leftward direction that initiates the left-right asymmetry in the developing embryo. <sup id="cite_ref-Horani_12-2" class="reference"><a href="#cite_note-Horani-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larsen_42-0" class="reference"><a href="#cite_note-Larsen-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>Motile, multiple, 9+0 cilia are found on the epithelial cells of the <a href="/wiki/Choroid_plexus" title="Choroid plexus">choroid plexus</a>. Cilia also can change structure when introduced to hot temperatures and become sharp. They are present in large numbers on each cell and move relatively slowly, making them intermediate between motile and primary cilia. In addition to 9+0 cilia that are mobile, there are also solitary 9+2 cilia that stay immobile found in hair cells.<sup id="cite_ref-Takeda_39-1" class="reference"><a href="#cite_note-Takeda-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Nodal_cilia">Nodal cilia</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=11" title="Edit section: Nodal cilia"><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:Nodal_cilia.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Nodal_cilia.jpg/220px-Nodal_cilia.jpg" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Nodal_cilia.jpg/330px-Nodal_cilia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Nodal_cilia.jpg/440px-Nodal_cilia.jpg 2x" data-file-width="560" data-file-height="372" /></a><figcaption><a href="/wiki/Scanning_electron_micrograph" class="mw-redirect" title="Scanning electron micrograph">Scanning electron micrograph</a> of nodal cilia on a <a href="/wiki/Mouse" title="Mouse">mouse</a> embryo</figcaption></figure> <p><b>Nodal cells</b> have a single cilium called a monocilium. They are present in the very early <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">development of the embryo</a> on the <a href="/wiki/Primitive_node" title="Primitive node">primitive node</a>. There are two areas of the node with different types of <b>nodal cilia</b>. On the central node are motile cilia, and on the peripheral area of the node the nodal cilia are modified motile.<sup id="cite_ref-Larsen_42-1" class="reference"><a href="#cite_note-Larsen-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>The motile cilia on the central cells rotate to generate the leftward flow of extracellular fluid needed to initiate the left-right asymmetry.<sup id="cite_ref-Larsen_42-2" class="reference"><a href="#cite_note-Larsen-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cilia_versus_flagella">Cilia versus flagella</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=12" title="Edit section: Cilia versus flagella"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Flagellum#Terminology" title="Flagellum">Flagellum § Terminology</a></div> <p>The motile cilia on <a href="/wiki/Sperm_cell" class="mw-redirect" title="Sperm cell">sperm cells</a> and many <a href="/wiki/Protozoan" class="mw-redirect" title="Protozoan">protozoans</a> enables swimming through liquids and are traditionally referred to as "<a href="/wiki/Flagella" class="mw-redirect" title="Flagella">flagella</a>".<sup id="cite_ref-Haimo_JCB198112_3-1" class="reference"><a href="#cite_note-Haimo_JCB198112-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> As these protrusions are structurally identical to motile cilia, attempts at preserving this terminology include making a distinction by morphology ("flagella" are typically longer than ordinary cilia and have a different undulating motion)<sup id="cite_ref-Alberts1_4-1" class="reference"><a href="#cite_note-Alberts1-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> and by number.<sup id="cite_ref-pmid20145000_43-0" class="reference"><a href="#cite_note-pmid20145000-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Microorganisms">Microorganisms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=13" title="Edit section: Microorganisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Ciliate" title="Ciliate">Ciliates</a> are <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a> that possess motile cilia exclusively and use them for either locomotion or to simply move liquid over their surface. A <i><a href="/wiki/Paramecium" title="Paramecium">Paramecium</a></i> for example is covered in thousands of cilia that enable its swimming. These motile cilia have been shown to be also sensory.<sup id="cite_ref-Valentine_44-0" class="reference"><a href="#cite_note-Valentine-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Ciliogenesis">Ciliogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=14" title="Edit section: Ciliogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Ciliogenesis" title="Ciliogenesis">Ciliogenesis</a></div> <p>Cilia are formed through the process of <a href="/wiki/Ciliogenesis" title="Ciliogenesis">ciliogenesis</a>. An early step is docking of the basal body to the growing ciliary membrane, after which the transition zone forms. The building blocks of the ciliary axoneme, such as <a href="/wiki/Tubulins" class="mw-redirect" title="Tubulins">tubulins</a>, are added at the ciliary tips through a process that depends partly on <a href="/wiki/Intraflagellar_transport" title="Intraflagellar transport">intraflagellar transport</a> (IFT).<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Exceptions include <i>Drosophila</i> sperm and <i><a href="/wiki/Plasmodium_falciparum" title="Plasmodium falciparum">Plasmodium falciparum</a></i> flagella formation, in which cilia assemble in the cytoplasm.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p>At the base of the cilium where it attaches to the cell body is the <a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">microtubule organizing center</a>, the <a href="/wiki/Basal_body" title="Basal body">basal body</a>. Some basal body proteins as <a href="/wiki/CEP164" title="CEP164">CEP164</a>, <a href="/wiki/ODF2" title="ODF2">ODF2</a><sup id="cite_ref-entrez2_48-0" class="reference"><a href="#cite_note-entrez2-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/CEP170" title="CEP170">CEP170</a>,<sup id="cite_ref-edoc_49-0" class="reference"><a href="#cite_note-edoc-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> are required for the formation and the stability of the cilium. </p><p>In effect, the cilium is a <a href="/wiki/Molecular_machine" title="Molecular machine">nanomachine</a> composed of perhaps over 600 proteins in molecular complexes, many of which also function independently as nanomachines. <a href="/wiki/Flexible_linker" class="mw-redirect" title="Flexible linker">Flexible linkers</a> allow the <a href="/wiki/Protein_domain#Domains_and_protein_flexibility" title="Protein domain">mobile protein domains</a> connected by them to recruit their binding partners and induce long-range <a href="/wiki/Allostery" class="mw-redirect" title="Allostery">allostery</a> via <a href="/wiki/Protein_dynamics#Global_flexibility:_multiple_domains" title="Protein dynamics">protein domain dynamics</a>.<sup id="cite_ref-Satir2008_35-1" class="reference"><a href="#cite_note-Satir2008-35"><span class="cite-bracket">[</span>35<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=Cilium&action=edit&section=15" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Dynein" title="Dynein">dynein</a> in the axoneme – <a href="/wiki/Axonemal_dynein" class="mw-redirect" title="Axonemal dynein">axonemal dynein</a> forms bridges between neighbouring microtubule doublets. When <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> activates the motor domain of dynein, it attempts to walk along the adjoining microtubule doublet. This would force the adjacent doublets to slide over one another if not for the presence of <a href="/wiki/Nexin" title="Nexin">nexin</a> between the microtubule doublets. And thus the force generated by dynein is instead converted into a bending motion.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-King_51-0" class="reference"><a href="#cite_note-King-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Sensing_the_extracellular_environment">Sensing the extracellular environment</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=16" title="Edit section: Sensing the extracellular environment"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some primary cilia on <a href="/wiki/Epithelial" class="mw-redirect" title="Epithelial">epithelial</a> cells in eukaryotes act as <i>cellular antennae</i>, providing <a href="/wiki/Molecular_sensor" title="Molecular sensor">chemosensation</a>, <a href="/wiki/Thermosensation" class="mw-redirect" title="Thermosensation">thermosensation</a> and <a href="/wiki/Mechanosensation" title="Mechanosensation">mechanosensation</a> of the extracellular environment.<sup id="cite_ref-Adams2008_52-0" class="reference"><a href="#cite_note-Adams2008-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Singla_629–633_10-2" class="reference"><a href="#cite_note-Singla_629–633-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> These cilia then play a role in mediating specific signalling cues, including soluble factors in the external cell environment, a <a href="/wiki/Secretion" title="Secretion">secretory</a> role in which a soluble protein is released to have an effect downstream of the fluid flow, and mediation of fluid flow if the cilia are <a href="/wiki/Motility" title="Motility">motile</a>.<sup id="cite_ref-Adams2008_52-1" class="reference"><a href="#cite_note-Adams2008-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>Some <a href="/wiki/Epithelial" class="mw-redirect" title="Epithelial">epithelial</a> cells are ciliated, and they commonly exist as a sheet of polarized cells forming a tube or tubule with cilia projecting into the <a href="/wiki/Lumen_(anatomy)" title="Lumen (anatomy)">lumen</a>. This sensory and signalling role puts cilia in a central role for maintaining the local cellular environment and may be why <a href="/wiki/Ciliopathy" title="Ciliopathy">ciliary defects</a> cause such a wide range of human diseases.<sup id="cite_ref-Reiter_533–547_31-1" class="reference"><a href="#cite_note-Reiter_533–547-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>In the embryo, <a href="/wiki/Nodal_cilia" class="mw-redirect" title="Nodal cilia">nodal cilia</a> are used to direct the flow of extracellular fluid. This leftward movement is to generate <a href="/wiki/Left-right_asymmetry" title="Left-right asymmetry">left-right asymmetry</a> across the midline of the embryo. Central cilia coordinate their rotational beating while the immotile cilia on the sides sense the direction of the flow.<sup id="cite_ref-Larsen_42-3" class="reference"><a href="#cite_note-Larsen-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Studies in mice suggest a biophysical mechanism by which the direction of flow is sensed.<sup id="cite_ref-science20230105_55-0" class="reference"><a href="#cite_note-science20230105-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Axo-ciliary_synapse">Axo-ciliary synapse</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=17" title="Edit section: Axo-ciliary synapse"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>With axo-ciliary <a href="/wiki/Synapse" title="Synapse">synapses</a>, there is communication between <a href="/wiki/Serotonin" title="Serotonin">serotonergic</a> <a href="/wiki/Axon" title="Axon">axons</a> and primary cilia of <a href="/wiki/Hippocampus_anatomy#Basic_hippocampal_circuit" title="Hippocampus anatomy">CA1</a> <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal</a> <a href="/wiki/Neuron" title="Neuron">neurons</a> that alters the neuron's <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> state in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> – "a way to change what is being transcribed or made in the nucleus" via this signalling distinct from that at the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> which also is longer-term.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </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=Cilium&action=edit&section=18" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Ciliopathy" title="Ciliopathy">Ciliopathy</a></div> <p>Ciliary defects can lead to a number of human diseases.<sup id="cite_ref-Reiter_533–547_31-2" class="reference"><a href="#cite_note-Reiter_533–547-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> Defects in cilia adversely affect many critical signaling pathways essential to embryonic development and to adult physiology, and thus offer a plausible hypothesis for the often <a href="/wiki/Pleiotropic" class="mw-redirect" title="Pleiotropic">multi-symptom</a> nature of diverse ciliopathies.<sup id="cite_ref-badano2006_30-1" class="reference"><a href="#cite_note-badano2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Reiter_533–547_31-3" class="reference"><a href="#cite_note-Reiter_533–547-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Known ciliopathies include <a href="/wiki/Primary_ciliary_dyskinesia" title="Primary ciliary dyskinesia">primary ciliary dyskinesia</a>, <a href="/wiki/Bardet%E2%80%93Biedl_syndrome" title="Bardet–Biedl syndrome">Bardet–Biedl syndrome</a>, <a href="/wiki/Polycystic_kidney_disease" title="Polycystic kidney disease">polycystic kidney</a> and <a href="/wiki/Polycystic_liver_disease" title="Polycystic liver disease">liver disease</a>, <a href="/wiki/Nephronophthisis" title="Nephronophthisis">nephronophthisis</a>, <a href="/wiki/Alstr%C3%B6m_syndrome" title="Alström syndrome">Alström syndrome</a>, <a href="/wiki/Meckel%E2%80%93Gruber_syndrome" title="Meckel–Gruber syndrome">Meckel–Gruber syndrome</a>, <a href="/wiki/Sensenbrenner_syndrome" title="Sensenbrenner syndrome">Sensenbrenner syndrome</a> and some forms of <a href="/wiki/Retinopathy" title="Retinopathy">retinal degeneration</a>.<sup id="cite_ref-badano2006_30-2" class="reference"><a href="#cite_note-badano2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Adams2008_52-2" class="reference"><a href="#cite_note-Adams2008-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Genetic mutations compromising the proper functioning of cilia, <a href="/wiki/Ciliopathy" title="Ciliopathy">ciliopathies</a>, can cause chronic disorders such as <a href="/wiki/Primary_ciliary_dyskinesia" title="Primary ciliary dyskinesia">primary ciliary dyskinesia</a> (PCD), <a href="/wiki/Nephronophthisis" title="Nephronophthisis">nephronophthisis</a>, and <a href="/wiki/Senior%E2%80%93L%C3%B8ken_syndrome" title="Senior–Løken syndrome">Senior–Løken syndrome</a>. In addition, a defect of the primary cilium in the <a href="/wiki/Nephron#Renal_tubule" title="Nephron">renal tubule</a> cells can lead to <a href="/wiki/Polycystic_kidney_disease" title="Polycystic kidney disease">polycystic kidney disease</a> (PKD). In another genetic disorder called <a href="/wiki/Bardet%E2%80%93Biedl_syndrome" title="Bardet–Biedl syndrome">Bardet–Biedl syndrome</a> (BBS), the mutant gene products are the components in the basal body and cilia.<sup id="cite_ref-badano2006_30-3" class="reference"><a href="#cite_note-badano2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Defects in cilia cells are linked to obesity and often pronounced in type 2 diabetes. Several studies already showed impaired glucose tolerance and reduction in the insulin secretion in the ciliopathy models. Moreover, the number and length of cilia was decreased in the <a href="/wiki/Type_2_diabetes" title="Type 2 diabetes">type 2 diabetes</a> models.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Epithelial_sodium_channels" class="mw-redirect" title="Epithelial sodium channels">Epithelial sodium channels</a> (ENaCs) that are expressed along the length of cilia regulate <a href="/wiki/Periciliary_liquid_layer" class="mw-redirect" title="Periciliary liquid layer">periciliary fluid</a> level. Mutations that decrease the activity of ENaCs result in multisystem <a href="/wiki/Pseudohypoaldosteronism" title="Pseudohypoaldosteronism">pseudohypoaldosteronism</a>, that is associated with fertility problems.<sup id="cite_ref-2012-Enuka_13-5" class="reference"><a href="#cite_note-2012-Enuka-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Cystic_fibrosis" title="Cystic fibrosis">cystic fibrosis</a> that results from mutations in the chloride channel <a href="/wiki/Cystic_fibrosis_transmembrane_conductance_regulator" title="Cystic fibrosis transmembrane conductance regulator">CFTR</a>, ENaC activity is enhanced leading to a severe reduction of the fluid level that causes complications and infections in the respiratory airways.<sup id="cite_ref-2016-Hanukoglu_41-1" class="reference"><a href="#cite_note-2016-Hanukoglu-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p><p>Since the flagellum of human sperm has the same internal structure of a cilium, ciliary dysfunction can also be responsible for male infertility.<sup id="cite_ref-pmid16850538_60-0" class="reference"><a href="#cite_note-pmid16850538-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p><p>There is an association of primary ciliary dyskinesia with left-right anatomic abnormalities such as <a href="/wiki/Situs_inversus" title="Situs inversus">situs inversus</a> (a combination of findings is known as <a href="/wiki/Kartagener_syndrome" class="mw-redirect" title="Kartagener syndrome">Kartagener syndrome</a>), and <a href="/wiki/Situs_ambiguus" title="Situs ambiguus">situs ambiguus</a> (also known as <i>Heterotaxy syndrome</i>).<sup id="cite_ref-Worsley_61-0" class="reference"><a href="#cite_note-Worsley-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> These left-right anatomic abnormalities can also result in <a href="/wiki/Congenital_heart_disease" class="mw-redirect" title="Congenital heart disease">congenital heart disease</a>.<sup id="cite_ref-pmid17515466_62-0" class="reference"><a href="#cite_note-pmid17515466-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> It has been shown that proper cilial function is responsible for the normal left-right asymmetry in mammals.<sup id="cite_ref-pmid12888012_63-0" class="reference"><a href="#cite_note-pmid12888012-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> </p><p>The diverse outcomes caused by ciliary dysfunction may result from alleles of different strengths that compromise ciliary functions in different ways or to different extents. Many ciliopathies are inherited in a Mendelian manner, but specific genetic interactions between distinct functional ciliary complexes, such as transition zone and BBS complexes, can alter the phenotypic manifestations of recessive ciliopathies.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Some mutations in transition zone proteins can cause specific serious ciliopathies.<sup id="cite_ref-Cavalier-Smith_66-0" class="reference"><a href="#cite_note-Cavalier-Smith-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Extracellular_changes">Extracellular changes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=19" title="Edit section: Extracellular changes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Reduction of cilia function can also result from infection. Research into <a href="/wiki/Biofilm" title="Biofilm">biofilms</a> has shown that bacteria can alter cilia. A biofilm is a community of bacteria of either the same or multiple species of bacteria. The cluster of cells secretes different factors which form an extracellular matrix. Cilia in the respiratory system is known to move mucus and pathogens out of the airways. It has been found that patients with biofilm positive infections have impaired cilia function. The impairment may present as decreased motion or reduction in the number of cilia. Though these changes result from an external source, they still effect the pathogenicity of the bacteria, progression of infection, and how it is treated.<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p><p>The transportation of the <a href="/wiki/Oocyte" title="Oocyte">immature egg cell</a>, and the embryo to the <a href="/wiki/Uterus" title="Uterus">uterus</a> for <a href="/wiki/Implantation_(human_embryo)" class="mw-redirect" title="Implantation (human embryo)">implantation</a> depends on the combination of regulated smooth muscle contractions, and ciliary beating. Dysfunction in this transportation can result in an <a href="/wiki/Ectopic_pregnancy" title="Ectopic pregnancy">ectopic pregnancy</a> where the embryo is implanted (usually) in the <a href="/wiki/Fallopian_tube" title="Fallopian tube">fallopian tube</a> before reaching its proper destination of the uterus. Many factors can affect this stage including infection and menstrual cycle hormones. Smoking (causing inflammation), and infection can reduce the numbers of cilia, and the ciliary beat can be affected by hormonal changes.<sup id="cite_ref-Panelli_15-2" class="reference"><a href="#cite_note-Panelli-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Primary_cilia_in_pancreatic_cells">Primary cilia in pancreatic cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cilium&action=edit&section=20" title="Edit section: Primary cilia in pancreatic cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The pancreas is a mixture of highly differentiated exocrine and endocrine cells. Primary cilia are present in exocrine cells, which are centroacinar duct cells.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hegyi_32-1" class="reference"><a href="#cite_note-Hegyi-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Endocrine tissue is composed of different hormone-secreting cells. Insulin-secreting beta cells and glucagon-secreting alpha cells are highly ciliated.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> </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=Cilium&action=edit&section=21" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Molecular_machine#Biological_molecular_machines" title="Molecular machine">Molecular machine#Biological molecular machines</a> – Molecular-scale artificial or biological device</li> <li><a href="/wiki/Protein_dynamics#Global_flexibility:_multiple_domains" title="Protein dynamics">Protein dynamics#Global flexibility: multiple domains</a> – Study of how proteins move and change shape</li> <li><a href="/wiki/Protein_domain#Domains_and_protein_flexibility" title="Protein domain">Protein domain#Domains and protein flexibility</a> – Self-stable region of a protein's chain that folds independently from the rest</li> <li><a href="/wiki/Stereocilia" title="Stereocilia">Stereocilia</a> – Cell structure</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=Cilium&action=edit&section=22" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-MW1-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-MW1_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.merriam-webster.com/dictionary/cilium">"Definition of CILIUM"</a>. <i>www.merriam-webster.com</i><span class="reference-accessdate">. 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href="https://web.archive.org/web/20060710113703/http://www.hhmi.org/bulletin/sept2005/pdf/Cilia.pdf">Brief summary of importance of cilia to many organs in human physiology</a></li> <li><a rel="nofollow" class="external text" href="http://www.ciliaproteome.org">The Ciliary Proteome Web Page at Johns Hopkins</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul 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.navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Structures_of_the_cell_/_organelles" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cellular_structures" title="Template:Cellular structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cellular_structures" title="Template talk:Cellular structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cellular_structures" title="Special:EditPage/Template:Cellular structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Structures_of_the_cell_/_organelles" style="font-size:114%;margin:0 4em">Structures of the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> / <a href="/wiki/Organelle" title="Organelle">organelles</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endomembrane_system" title="Endomembrane system">Endomembrane <br />system</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/Cell_membrane" title="Cell membrane">Cell membrane</a></li> <li><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></li> <li><a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">Endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a></li> <li><a href="/wiki/Parenthesome" title="Parenthesome">Parenthesome</a></li> <li><a href="/wiki/Autophagosome" title="Autophagosome">Autophagosome</a></li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a> <ul><li><a href="/wiki/Exosome_(vesicle)" title="Exosome (vesicle)">Exosome</a></li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Endosome" title="Endosome">Endosome</a></li> <li><a href="/wiki/Phagosome" title="Phagosome">Phagosome</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Acrosome" title="Acrosome">Acrosome</a></li></ul></li> <li><a href="/wiki/Granule_(cell_biology)" title="Granule (cell biology)">Cytoplasmic granule</a> <ul><li><a href="/wiki/Melanosome" title="Melanosome">Melanosome</a></li> <li><a href="/wiki/Microbody" title="Microbody">Microbody</a></li> <li><a href="/wiki/Glyoxysome" title="Glyoxysome">Glyoxysome</a></li> <li><a href="/wiki/Peroxisome" title="Peroxisome">Peroxisome</a></li> <li><a href="/wiki/Weibel%E2%80%93Palade_body" title="Weibel–Palade body">Weibel–Palade body</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</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/Microfilament" title="Microfilament">Microfilament</a></li> <li><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate filament</a></li> <li><a href="/wiki/Microtubule" title="Microtubule">Microtubule</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a></li> <li><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organizing center</a> <ul><li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Centriole" title="Centriole">Centriole</a></li> <li><a href="/wiki/Basal_body" title="Basal body">Basal body</a></li> <li><a href="/wiki/Spindle_pole_body" title="Spindle pole body">Spindle pole body</a></li></ul></li> <li><a href="/wiki/Myofibril" title="Myofibril">Myofibril</a></li> <li><a href="/wiki/Undulipodium" title="Undulipodium">Undulipodium</a> <ul><li><a class="mw-selflink selflink">Cilium</a></li> <li><a href="/wiki/Flagellum#Eukaryotic" title="Flagellum">Flagellum</a></li> <li><a href="/wiki/Axoneme" title="Axoneme">Axoneme</a></li> <li><a href="/wiki/Radial_spoke" title="Radial spoke">Radial spoke</a></li></ul></li> <li><a href="/wiki/Pseudopodium" class="mw-redirect" title="Pseudopodium">Pseudopodium</a> <ul><li><a href="/wiki/Lamellipodium" title="Lamellipodium">Lamellipodium</a></li> <li><a href="/wiki/Filopodium" class="mw-redirect" title="Filopodium">Filopodium</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbionts</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/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Amyloplast" title="Amyloplast">Amyloplast</a></li> <li><a href="/wiki/Elaioplast" title="Elaioplast">Elaioplast</a></li> <li><a href="/wiki/Proteinoplast" title="Proteinoplast">Proteinoplast</a></li> <li><a href="/wiki/Tannosome" title="Tannosome">Tannosome</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other internal</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/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Spliceosome" title="Spliceosome">Spliceosome</a></li> <li><a href="/wiki/Vault_(organelle)" title="Vault (organelle)">Vault</a></li></ul></li> <li><a href="/wiki/Cytoplasm" title="Cytoplasm">Cytoplasm</a> <ul><li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a></li> <li><a href="/wiki/Cytoplasmic_inclusion" class="mw-redirect" title="Cytoplasmic inclusion">Inclusions</a></li></ul></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</a></li> <li><a href="/wiki/Magnetosome" title="Magnetosome">Magnetosome</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">External</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/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Extracellular_matrix" title="Extracellular matrix">Extracellular matrix</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="Proteins_of_epithelium" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template: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 class="mw-selflink selflink">Cilia</a>/<a href="/wiki/Kinocilium" title="Kinocilium">Kinocilium</a></li> <li><a href="/wiki/Microvillus" title="Microvillus">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="Microbiology:_Protistology:_Protists" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Protist_structures" title="Template:Protist structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Protist_structures" title="Template talk:Protist structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Protist_structures" title="Special:EditPage/Template:Protist structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Microbiology:_Protistology:_Protists" style="font-size:114%;margin:0 4em"><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a>: <a href="/wiki/Protistology" title="Protistology">Protistology</a>: <a href="/wiki/Protist" title="Protist">Protists</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Former<br />classifications</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/Protozoa" title="Protozoa">Protozoa</a> <ul><li><a href="/wiki/Flagellate" title="Flagellate">Mastigophora/Flagellates</a></li> <li><a href="/wiki/Amoeba" title="Amoeba">Sarcodina/Amoeboids</a> <ul><li><a href="/wiki/Testate_amoebae" title="Testate amoebae">Testate</a></li> <li><a href="/wiki/Heliozoa" title="Heliozoa">Heliozoa</a></li></ul></li> <li><a href="/wiki/Infusoria" title="Infusoria">Infusoria</a>/<a href="/wiki/Ciliate" title="Ciliate">Ciliates</a></li> <li><a href="/wiki/Apicomplexa#Taxonomy" title="Apicomplexa">Sporozoa</a></li></ul></li> <li><a href="/wiki/Algae" title="Algae">Algae</a></li> <li><a href="/wiki/Cryptogam" title="Cryptogam">Cryptogams</a></li> <li><a href="/wiki/Thallophyte" title="Thallophyte">Thallophytes</a></li> <li><a href="/wiki/Fungus#Fungus-like_organisms" title="Fungus">Fungus-like organisms</a></li> <li><a href="/wiki/Slime_mold" title="Slime mold">Slime molds</a></li> <li><a href="/wiki/Nomenclature_codes#Ambiregnal_protists" title="Nomenclature codes">Ambiregnal protists</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Morphology</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%;background:lightgreen;"><a href="/wiki/Archaeplastida" title="Archaeplastida">Archaeplastida</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background:lightgreen;"><span class="nobold"><a href="/wiki/Viridiplantae" title="Viridiplantae">Chloroplastida</a></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>"<a href="/wiki/Green_algae" title="Green algae">green algae</a>": <a href="/wiki/Phycoplast" title="Phycoplast">Phycoplast</a></li> <li><a href="/wiki/Phragmoplast" title="Phragmoplast">Phragmoplast</a></li> <li><a href="/wiki/Flagellum#Types" title="Flagellum">Flagellar apparatus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:lightgreen;"><span class="nobold"><a href="/wiki/Glaucophyte" title="Glaucophyte">Glaucophytes</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cyanelle" class="mw-redirect" title="Cyanelle">Cyanelles</a></li> <li><a href="/wiki/Phycobilisome" title="Phycobilisome">Phycobilisomes</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:lightgreen;"><span class="nobold"><a href="/wiki/Red_algae" title="Red algae">Red algae</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pit_connection" title="Pit connection">Pit connection</a></li> <li><a href="/wiki/Phycobilisome" title="Phycobilisome">Phycobilisomes</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><a href="/wiki/Hacrobia" title="Hacrobia">Hacrobia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Cryptomonad" title="Cryptomonad">Cryptophyta</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mastigoneme" title="Mastigoneme">Mastigoneme</a></li> <li><a href="/wiki/Periplast" title="Periplast">Periplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Haptophyte" title="Haptophyte">Haptophyte</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Coccolith" title="Coccolith">Coccolith</a></li> <li><a href="/wiki/Haptophyte#Characteristics" title="Haptophyte">Haptonema</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><a href="/wiki/Stramenopile" title="Stramenopile">Stramenopiles</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold">General:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mastigoneme" title="Mastigoneme">Mastigoneme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Diatom" title="Diatom">Diatoms</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Frustule" title="Frustule">Frustule</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Brown_algae" title="Brown algae">Brown algae</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lamina_(algae)" title="Lamina (algae)">Lamina</a></li> <li><a href="/wiki/Pneumatocyst" title="Pneumatocyst">Pneumatocyst</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><a href="/wiki/Alveolate" title="Alveolate">Alveolata</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold">General:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alveolate#Characteristics" title="Alveolate"> Alveoli</a></li> <li><a href="/wiki/Trichocyst" title="Trichocyst">Trichocyst</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Dinoflagellate" title="Dinoflagellate">Dinoflagellates</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dinokaryon" title="Dinokaryon">Dinokaryon</a></li> <li><a href="/wiki/Dinocyst" title="Dinocyst">Dinocyst</a></li> <li><a href="/wiki/Theca" title="Theca">Theca</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Ciliate" title="Ciliate">Ciliates</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Cilium</a></li> <li><a href="/wiki/Cirrus_(biology)" title="Cirrus (biology)">Cirrus</a></li> <li><a href="/wiki/Macronucleus" title="Macronucleus">Macronucleus</a></li> <li><a href="/wiki/Micronucleus" title="Micronucleus">Micronucleus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#ADFF2F;"><span class="nobold"><a href="/wiki/Apicomplexa" title="Apicomplexa">Apicomplexans</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Rhoptry" title="Rhoptry">Rhoptry</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li> <li><a href="/wiki/Microneme" title="Microneme">Microneme</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:rgb(225,204,252);"><a href="/wiki/Rhizaria" title="Rhizaria">Rhizaria</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Phytomyxea" scope="row" class="navbox-group" style="width:1%;background:rgb(225,204,252);"><span class="nobold"><a href="/wiki/Phytomyxea" title="Phytomyxea">Phytomyxea</a></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nuclear_division" class="mw-redirect" title="Nuclear division">Cruciform division</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#F0E68C;"><a href="/wiki/Excavata" title="Excavata">Excavate</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background:#F0E68C;"><span class="nobold"><a href="/wiki/Kinetoplastida" title="Kinetoplastida">Kinetoplastids</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Kinetoplast" title="Kinetoplast">Kinetoplast</a></li> <li><a href="/wiki/Glycosome" title="Glycosome">Glycosome</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#F0E68C;"><span class="nobold"><a href="/wiki/Euglenid" title="Euglenid">Euglenoidea</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Periplast" title="Periplast">Periplast/pellicle</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#FFC8A0;"><a href="/wiki/Amoebozoa" title="Amoebozoa">Amoebozoa</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Dictyostelids:" scope="row" class="navbox-group" style="width:1%;background:#FFC8A0;"><span class="nobold"><a href="/wiki/Dictyostelid" title="Dictyostelid">Dictyostelids</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Macrocyst" title="Macrocyst">Macrocyst</a></li> <li><a href="/wiki/Sorocarp" title="Sorocarp">Sorocarp</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/Opisthokont" title="Opisthokont">Opisthokonta</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Choanoflagellates:" scope="row" class="navbox-group" style="width:1%;background:#D3D3A4;"><span class="nobold"><a href="/wiki/Choanoflagellate" title="Choanoflagellate">Choanoflagellates</a>:</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Microvillus" title="Microvillus">Collar of microvilli</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="https://de.wikipedia.org/wiki/Organisationsstufe" class="extiw" title="de:Organisationsstufe">Levels of<br />organization</a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Unicellular" class="mw-redirect" title="Unicellular">Unicellular</a> <ul><li><a href="/wiki/Flagellate" title="Flagellate">Monadoid</a></li> <li><a href="/wiki/Amoeboid" class="mw-redirect" title="Amoeboid">Amoeboid</a></li> <li><a href="/wiki/Coccoid_(disambiguation)" class="mw-redirect mw-disambig" title="Coccoid (disambiguation)">Coccoid</a></li> <li><a href="/wiki/Amoeboflagellate" title="Amoeboflagellate">Amoeboflagellate</a></li></ul></li> <li><a href="/wiki/Colony_(biology)" title="Colony (biology)">Colonial</a> <a href="/wiki/Sensu_stricto" class="mw-redirect" title="Sensu stricto">s.s.</a> <ul><li>Colonial flagellated</li> <li>Tetrasporal/capsal/palmelloid</li> <li>Sarcinoid</li> <li><a href="/wiki/Coenobium_(morphology)" class="mw-redirect" title="Coenobium (morphology)">Coenobial</a></li></ul></li> <li>Filamentous/trichal/<a href="/wiki/Hyphal" class="mw-redirect" title="Hyphal">hyphal</a></li> <li><a href="/wiki/Parenchymatous" class="mw-redirect" title="Parenchymatous">Parenchymatous</a></li> <li><a href="/wiki/Pseudoparenchyma" class="mw-redirect" title="Pseudoparenchyma">Pseudoparenchymatous</a>/plektenchymatic</li> <li>Membranous/thalloid/foliaceous</li> <li><a href="/wiki/Multinucleated" class="mw-redirect" title="Multinucleated">Multinucleated</a> <ul><li><a href="/wiki/Syncytial" class="mw-redirect" title="Syncytial">Syncytial</a></li> <li><a href="/wiki/Coenocytic" class="mw-redirect" title="Coenocytic">Coenocytic</a> <ul><li>Siphonous</li> <li>Siphonocladous</li></ul></li></ul></li> <li><a href="/wiki/Multicellular" class="mw-redirect" title="Multicellular">Multicellular</a> <a href="/wiki/Sensu_stricto" class="mw-redirect" title="Sensu stricto">s.s.</a>/<a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissular</a>/histonal</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Cell_wall#Other_cell_coverings" title="Cell wall">Cell surface<br />structures</a></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_membrane" title="Cell membrane">Simple cell membrane</a></li> <li><a href="/wiki/Mucilage" title="Mucilage">Mucilage</a></li> <li><a href="/wiki/Scale_(anatomy)" class="mw-redirect" title="Scale (anatomy)">Scale</a></li> <li><a href="/wiki/Frustule" title="Frustule">Frustule</a></li> <li><a href="/wiki/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Lorica_(biology)" title="Lorica (biology)">Lorica</a></li> <li><a href="/wiki/Skeleton" title="Skeleton">Skeleton</a></li> <li><a href="/wiki/Test_(biology)" title="Test (biology)">Test</a></li> <li><a href="/wiki/Theca" title="Theca">Theca</a></li> <li><a href="/wiki/Periplast" title="Periplast">Periplast/pellicle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold">Locomotion</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Flagellum" title="Flagellum">Flagellum</a></li> <li><a class="mw-selflink selflink">Cilium</a></li> <li><a href="/wiki/Pseudopodia" title="Pseudopodia">Pseudopodia</a></li> <li><a href="/wiki/Gliding_motility" title="Gliding motility">Gliding motility</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold">Mitochondria</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li> <li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold">Nucleus</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nucleomorph" title="Nucleomorph">Nucleomorph</a></li> <li><a href="/wiki/Multinucleate" title="Multinucleate">Multinucleate cells</a> <ul><li><a href="/wiki/Plasmodium_(life_cycle)" title="Plasmodium (life cycle)">Plasmodium</a></li></ul></li> <li><a href="/wiki/Dikaryon" title="Dikaryon">Dikaryon</a></li> <li><a href="/wiki/Heterokaryon" title="Heterokaryon">Heterokaryon</a></li> <li><a href="/wiki/Mitosis" title="Mitosis">Mitosis in protists</a> <ul><li>Open</li> <li>Closed</li> <li>Orthomitosis</li> <li>Pleuromitosis</li></ul></li> <li><a href="/wiki/Meiosis#Occurrence_in_eukaryotic_life_cycles" title="Meiosis">Meiosis in protists</a> <ul><li>Gametic</li> <li>Zygotic</li> <li>Sporic</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold">Other</span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Microbial_cyst" title="Microbial cyst">Cyst</a></li> <li><a href="/wiki/Cytostome" title="Cytostome">Cytostome</a></li> <li><a href="/wiki/Fimbria_(bacteriology)" class="mw-redirect" title="Fimbria (bacteriology)">Fimbriae</a></li> <li><a href="/wiki/Extrusome" title="Extrusome">Extrusome</a></li> <li><a href="/wiki/Contractile_vacuole" title="Contractile vacuole">Contractile vacuole</a></li> <li><a href="/wiki/Eyespot_apparatus" title="Eyespot apparatus">Eyespot apparatus</a></li> <li><a href="/wiki/Pyrenoid" title="Pyrenoid">Pyrenoid</a></li> <li><a href="/wiki/Axostyle" title="Axostyle">Axostyle</a></li> <li><a href="/wiki/Mastigont_system" title="Mastigont system">Mastigont system</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%">Ecology and<br />physiology</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/Microbial_ecology" title="Microbial ecology">Microbial ecology</a> <ul><li><a href="/wiki/Microbial_biogeography" title="Microbial biogeography">Microbial biogeography</a></li> <li><a href="/wiki/Lourens_Baas_Becking#Baas_Becking_hypothesis" title="Lourens Baas Becking">Baas-Becking hypothesis</a></li></ul></li> <li><a href="/wiki/Developmental_biology" title="Developmental biology">Development</a>/<a href="/wiki/Reproduction" title="Reproduction">Reproduction</a> <ul><li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles</a></li> <li><a href="/wiki/Fertilization#Fertilization_in_protists" class="mw-redirect" title="Fertilization">Fertilization</a></li></ul></li> <li>Nutrition: <a href="/wiki/Autotrophy" class="mw-redirect" title="Autotrophy">Autotrophy</a> <ul><li><a href="/wiki/Phototrophy" class="mw-redirect" title="Phototrophy">Phototrophy</a></li></ul></li> <li><a href="/wiki/Heterotrophy" class="mw-redirect" title="Heterotrophy">Heterotrophy</a> <ul><li><a href="/wiki/Phagotrophy" class="mw-redirect" title="Phagotrophy">Phagotrophy</a></li> <li><a href="/wiki/Osmotrophy" title="Osmotrophy">Osmotrophy</a></li> <li><a href="/wiki/Saprotrophic_nutrition" title="Saprotrophic nutrition">Saprotrophy</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a> <ul><li><a href="/wiki/Biotrophy" class="mw-redirect" title="Biotrophy">Biotrophy</a></li> <li><a href="/wiki/Necrotrophy" class="mw-redirect" title="Necrotrophy">Necrotrophy</a></li></ul></li></ul></li> <li><a href="/wiki/Mixotrophy" class="mw-redirect" title="Mixotrophy">Mixotrophy</a></li> <li><a href="/wiki/Auxotrophy" title="Auxotrophy">Auxotrophy</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link 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