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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Eukaryotic_cytoskeleton" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Eukaryotic_cytoskeleton"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Eukaryotic cytoskeleton</span> </div> </a> <button aria-controls="toc-Eukaryotic_cytoskeleton-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 Eukaryotic cytoskeleton subsection</span> </button> <ul id="toc-Eukaryotic_cytoskeleton-sublist" class="vector-toc-list"> <li id="toc-Microfilaments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Microfilaments"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Microfilaments</span> </div> </a> <ul id="toc-Microfilaments-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Intermediate_filaments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Intermediate_filaments"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Intermediate filaments</span> </div> </a> <ul id="toc-Intermediate_filaments-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Microtubules" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Microtubules"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Microtubules</span> </div> </a> <ul id="toc-Microtubules-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Comparison" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Comparison"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Comparison</span> </div> </a> <ul id="toc-Comparison-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Septins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Septins"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Septins</span> </div> </a> <ul id="toc-Septins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Spectrin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Spectrin"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.6</span> <span>Spectrin</span> </div> </a> <ul id="toc-Spectrin-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Yeast_cytoskeleton" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Yeast_cytoskeleton"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.7</span> <span>Yeast cytoskeleton</span> </div> </a> <ul id="toc-Yeast_cytoskeleton-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Prokaryotic_cytoskeleton" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Prokaryotic_cytoskeleton"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Prokaryotic cytoskeleton</span> </div> </a> <button aria-controls="toc-Prokaryotic_cytoskeleton-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 Prokaryotic cytoskeleton subsection</span> </button> <ul id="toc-Prokaryotic_cytoskeleton-sublist" class="vector-toc-list"> <li id="toc-FtsZ" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#FtsZ"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>FtsZ</span> </div> </a> <ul id="toc-FtsZ-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-MreB_and_ParM" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#MreB_and_ParM"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>MreB and ParM</span> </div> </a> <ul id="toc-MreB_and_ParM-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Crescentin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Crescentin"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Crescentin</span> </div> </a> <ul id="toc-Crescentin-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-The_cytoskeleton_and_cell_mechanics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#The_cytoskeleton_and_cell_mechanics"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>The cytoskeleton and cell mechanics</span> </div> </a> <ul id="toc-The_cytoskeleton_and_cell_mechanics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Long-range_order" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Long-range_order"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Long-range order</span> </div> </a> <ul id="toc-Long-range_order-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Common_features_and_differences_between_prokaryotes_and_eukaryotes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Common_features_and_differences_between_prokaryotes_and_eukaryotes"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Common features and differences between prokaryotes and eukaryotes</span> </div> </a> <ul id="toc-Common_features_and_differences_between_prokaryotes_and_eukaryotes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cytoplasmic_streaming" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Cytoplasmic_streaming"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Cytoplasmic streaming</span> </div> </a> <ul id="toc-Cytoplasmic_streaming-sublist" class="vector-toc-list"> </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">8</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">9</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">10</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">Cytoskeleton</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 66 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-66" 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">66 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%D9%8A%D9%83%D9%84_%D8%AE%D9%84%D9%88%D9%8A" title="هيكل خلوي – Arabic" lang="ar" hreflang="ar" data-title="هيكل خلوي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%95%E0%A7%8B%E0%A6%B7%E0%A6%95%E0%A6%99%E0%A7%8D%E0%A6%95%E0%A6%BE%E0%A6%B2" title="কোষকঙ্কাল – Bangla" lang="bn" hreflang="bn" data-title="কোষকঙ্কাল" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A6%D0%B8%D1%82%D0%BE%D1%81%D0%BA%D0%B5%D0%BB%D0%B5%D1%82" title="Цитоскелет – Bulgarian" lang="bg" hreflang="bg" data-title="Цитоскелет" 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/Citoskelet" title="Citoskelet – Bosnian" lang="bs" hreflang="bs" data-title="Citoskelet" 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/Citoesquelet" title="Citoesquelet – Catalan" lang="ca" hreflang="ca" data-title="Citoesquelet" 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/Cytoskelet" title="Cytoskelet – Czech" lang="cs" hreflang="cs" data-title="Cytoskelet" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Cytosgerbwd" title="Cytosgerbwd – Welsh" lang="cy" hreflang="cy" data-title="Cytosgerbwd" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target"><span>Cymraeg</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Cytoskelet" title="Cytoskelet – Danish" lang="da" hreflang="da" data-title="Cytoskelet" 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/Cytoskelett" title="Cytoskelett – German" lang="de" hreflang="de" data-title="Cytoskelett" 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/Ts%C3%BCtoskelett" title="Tsütoskelett – Estonian" lang="et" hreflang="et" data-title="Tsütoskelett" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CF%85%CF%84%CF%84%CE%B1%CF%81%CE%B9%CE%BA%CF%8C%CF%82_%CF%83%CE%BA%CE%B5%CE%BB%CE%B5%CF%84%CF%8C%CF%82" title="Κυτταρικός σκελετός – Greek" lang="el" hreflang="el" data-title="Κυτταρικός σκελετός" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Citoesqueleto" title="Citoesqueleto – Spanish" lang="es" hreflang="es" data-title="Citoesqueleto" 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/%C4%88elskeleto" title="Ĉelskeleto – Esperanto" lang="eo" hreflang="eo" data-title="Ĉelskeleto" 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/Zitoeskeleto" title="Zitoeskeleto – Basque" lang="eu" hreflang="eu" data-title="Zitoeskeleto" 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/%D8%A7%D8%B3%DA%A9%D9%84%D8%AA_%D8%B3%D9%84%D9%88%D9%84%DB%8C" title="اسکلت سلولی – Persian" lang="fa" hreflang="fa" data-title="اسکلت سلولی" 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/Cytosquelette" title="Cytosquelette – French" lang="fr" hreflang="fr" data-title="Cytosquelette" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/C%C3%ADtichreatlach" title="Cítichreatlach – Irish" lang="ga" hreflang="ga" data-title="Cítichreatlach" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gv mw-list-item"><a href="https://gv.wikipedia.org/wiki/Ushylagh_killag" title="Ushylagh killag – Manx" lang="gv" hreflang="gv" data-title="Ushylagh killag" data-language-autonym="Gaelg" data-language-local-name="Manx" class="interlanguage-link-target"><span>Gaelg</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Citoesqueleto" title="Citoesqueleto – Galician" lang="gl" hreflang="gl" data-title="Citoesqueleto" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%84%B8%ED%8F%AC%EA%B3%A8%EA%B2%A9" title="세포골격 – Korean" lang="ko" hreflang="ko" data-title="세포골격" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D5%BB%D5%BB%D5%A1%D5%AF%D5%B4%D5%A1%D5%AD%D6%84" title="Բջջակմախք – Armenian" lang="hy" hreflang="hy" data-title="Բջջակմախք" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%95%E0%A5%8B%E0%A4%B6%E0%A4%BF%E0%A4%95%E0%A4%BE_%E0%A4%95%E0%A4%82%E0%A4%95%E0%A4%BE%E0%A4%B2" title="कोशिका कंकाल – Hindi" lang="hi" hreflang="hi" data-title="कोशिका कंकाल" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Citoskelet" title="Citoskelet – Croatian" lang="hr" hreflang="hr" data-title="Citoskelet" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Sitoskeleton" title="Sitoskeleton – Indonesian" lang="id" hreflang="id" data-title="Sitoskeleton" 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-zu mw-list-item"><a href="https://zu.wikipedia.org/wiki/Isixhakangqamu" title="Isixhakangqamu – Zulu" lang="zu" hreflang="zu" data-title="Isixhakangqamu" data-language-autonym="IsiZulu" data-language-local-name="Zulu" class="interlanguage-link-target"><span>IsiZulu</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Frymisgrind" title="Frymisgrind – Icelandic" lang="is" hreflang="is" data-title="Frymisgrind" 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/Citoscheletro" title="Citoscheletro – Italian" lang="it" hreflang="it" data-title="Citoscheletro" 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%A9%D7%9C%D7%93_%D7%94%D7%AA%D7%90" title="שלד התא – Hebrew" lang="he" hreflang="he" data-title="שלד התא" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%95%E0%B3%8B%E0%B2%B6_%E0%B2%95%E0%B2%82%E0%B2%95%E0%B2%BE%E0%B2%B2" title="ಕೋಶ ಕಂಕಾಲ – Kannada" lang="kn" hreflang="kn" data-title="ಕೋಶ ಕಂಕಾಲ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%96%D0%B0%D1%81%D1%83%D1%88%D0%B0_%D2%9B%D0%B0%D2%A3%D2%9B%D0%B0%D1%81%D1%8B" title="Жасуша қаңқасы – Kazakh" lang="kk" hreflang="kk" data-title="Жасуша қаңқасы" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Peyker%C3%AA_xaney%C3%AA" title="Peykerê xaneyê – Kurdish" lang="ku" hreflang="ku" data-title="Peykerê xaneyê" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Cytosceletus" title="Cytosceletus – Latin" lang="la" hreflang="la" data-title="Cytosceletus" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Citoskelets" title="Citoskelets – Latvian" lang="lv" hreflang="lv" data-title="Citoskelets" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Zytoskelett" title="Zytoskelett – Luxembourgish" lang="lb" hreflang="lb" data-title="Zytoskelett" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target"><span>Lëtzebuergesch</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Citoskeletas" title="Citoskeletas – Lithuanian" lang="lt" hreflang="lt" data-title="Citoskeletas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Sitosceleto" title="Sitosceleto – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Sitosceleto" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target"><span>Lingua Franca Nova</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Citoszkeleton" title="Citoszkeleton – Hungarian" lang="hu" hreflang="hu" data-title="Citoszkeleton" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%A6%D0%B8%D1%82%D0%BE%D1%81%D0%BA%D0%B5%D0%BB%D0%B5%D1%82" title="Цитоскелет – Macedonian" lang="mk" hreflang="mk" data-title="Цитоскелет" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Cytoskelet" title="Cytoskelet – Dutch" lang="nl" hreflang="nl" data-title="Cytoskelet" 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%B4%B0%E8%83%9E%E9%AA%A8%E6%A0%BC" title="細胞骨格 – Japanese" lang="ja" hreflang="ja" data-title="細胞骨格" 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/Cytoskjelett" title="Cytoskjelett – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Cytoskjelett" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Citoesquel%C3%A8t" title="Citoesquelèt – Occitan" lang="oc" hreflang="oc" data-title="Citoesquelèt" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Cytoszkielet" title="Cytoszkielet – Polish" lang="pl" hreflang="pl" data-title="Cytoszkielet" 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/Citoesqueleto" title="Citoesqueleto – Portuguese" lang="pt" hreflang="pt" data-title="Citoesqueleto" 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/Citoschelet" title="Citoschelet – Romanian" lang="ro" hreflang="ro" data-title="Citoschelet" 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%A6%D0%B8%D1%82%D0%BE%D1%81%D0%BA%D0%B5%D0%BB%D0%B5%D1%82" title="Цитоскелет – Russian" lang="ru" hreflang="ru" data-title="Цитоскелет" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B7%83%E0%B7%99%E0%B7%99%E0%B6%BD_%E0%B7%83%E0%B7%90%E0%B6%9A%E0%B7%92%E0%B6%BD%E0%B7%8A%E0%B6%BD" title="සෙෙල සැකිල්ල – Sinhala" lang="si" hreflang="si" data-title="සෙෙල සැකිල්ල" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cytoskeleton" title="Cytoskeleton – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cytoskeleton" 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-sd mw-list-item"><a href="https://sd.wikipedia.org/wiki/%D8%B3%D8%A7%D8%A6%D9%BD%D9%88%D8%B3%DA%AA%D9%84%D9%BD%D9%86" title="سائٽوسڪلٽن – Sindhi" lang="sd" hreflang="sd" data-title="سائٽوسڪلٽن" data-language-autonym="سنڌي" data-language-local-name="Sindhi" class="interlanguage-link-target"><span>سنڌي</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Cytoskelet" title="Cytoskelet – Slovak" lang="sk" hreflang="sk" data-title="Cytoskelet" 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/Citoskelet" title="Citoskelet – Slovenian" lang="sl" hreflang="sl" data-title="Citoskelet" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%BE%DB%95%DB%8C%DA%A9%DB%95%D8%B1%DB%8C_%D8%AE%D8%A7%D9%86%DB%95" title="پەیکەری خانە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="پەیکەری خانە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A6%D0%B8%D1%82%D0%BE%D1%81%D0%BA%D0%B5%D0%BB%D0%B5%D1%82" title="Цитоскелет – Serbian" lang="sr" hreflang="sr" data-title="Цитоскелет" 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/Citoskelet" title="Citoskelet – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Citoskelet" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Solun_tukiranka" title="Solun tukiranka – Finnish" lang="fi" hreflang="fi" data-title="Solun tukiranka" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Cellskelett" title="Cellskelett – Swedish" lang="sv" hreflang="sv" data-title="Cellskelett" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Cytoskeleton" title="Cytoskeleton – Tagalog" lang="tl" hreflang="tl" data-title="Cytoskeleton" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%95%E0%AF%81%E0%AE%B4%E0%AE%BF%E0%AE%AF_%E0%AE%B5%E0%AE%A9%E0%AF%8D%E0%AE%95%E0%AF%82%E0%AE%9F%E0%AF%81" title="குழிய வன்கூடு – Tamil" lang="ta" hreflang="ta" data-title="குழிய வன்கூடு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%84%E0%B8%8B%E0%B9%82%E0%B8%97%E0%B8%AA%E0%B9%80%E0%B8%81%E0%B9%80%E0%B8%A5%E0%B8%95%E0%B8%B1%E0%B8%99" title="ไซโทสเกเลตัน – Thai" lang="th" hreflang="th" data-title="ไซโทสเกเลตัน" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/H%C3%BCcre_iskeleti" title="Hücre iskeleti – Turkish" lang="tr" hreflang="tr" data-title="Hücre iskeleti" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A6%D0%B8%D1%82%D0%BE%D1%81%D0%BA%D0%B5%D0%BB%D0%B5%D1%82" title="Цитоскелет – Ukrainian" lang="uk" hreflang="uk" data-title="Цитоскелет" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B3%D8%A7%D8%A6%D9%B9%D9%88%D8%B3%DA%A9%D9%84%D9%B9%D9%86" title="سائٹوسکلٹن – Urdu" lang="ur" hreflang="ur" data-title="سائٹوسکلٹن" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/B%E1%BB%99_x%C6%B0%C6%A1ng_t%E1%BA%BF_b%C3%A0o" title="Bộ xương tế bào – Vietnamese" lang="vi" hreflang="vi" data-title="Bộ xương tế bào" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E7%BB%86%E8%83%9E%E9%AA%A8%E6%9E%B6" title="细胞骨架 – Wu" lang="wuu" hreflang="wuu" data-title="细胞骨架" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E9%AA%A8%E6%A0%BC" title="細胞骨格 – Cantonese" lang="yue" hreflang="yue" data-title="細胞骨格" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%BB%86%E8%83%9E%E9%AA%A8%E6%9E%B6" title="细胞骨架 – Chinese" lang="zh" hreflang="zh" data-title="细胞骨架" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q154626#sitelinks-wikipedia" title="Edit interlanguage links" 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id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Network of filamentous proteins that forms the internal framework of cells</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox" style="width: 300px; padding: 0; border-collapse:collapse; background:transparent;"><tbody><tr><th colspan="2" class="infobox-above" style="background: rgba(0,0,0,0.1); padding: 5px; width: 290px;"><a href="/wiki/Cell_biology" title="Cell biology"><span style="color:#000;">Cell biology</span></a></th></tr><tr><td colspan="2" class="infobox-subheader" style="font-weight:bold; font-size:125%; background-color: #0073CF; color: #FFF;"><a href="/wiki/Animal_cell" class="mw-redirect" title="Animal cell"><span style="color: #FFF;">Animal cell diagram</span></a></td></tr><tr><td colspan="2" class="infobox-image" style="padding: 0;"><span typeof="mw:File"><a href="/wiki/File:Animal_Cell.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/300px-Animal_Cell.svg.png" decoding="async" width="300" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/450px-Animal_Cell.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/600px-Animal_Cell.svg.png 2x" data-file-width="1405" data-file-height="811" /></a></span><div class="infobox-caption" style="text-align:left; padding: 10px; background: rgba(0,0,0,0.02);">Components of a typical animal cell: <ol><li><a href="/wiki/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></li> <li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a> (dots as part of 5)</li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a></li> <li><a href="/wiki/Rough_endoplasmic_reticulum" class="mw-redirect" title="Rough endoplasmic reticulum">Rough endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> (or, Golgi body)</li> <li><a class="mw-selflink selflink">Cytoskeleton</a></li> <li><a href="/wiki/Smooth_endoplasmic_reticulum" class="mw-redirect" title="Smooth endoplasmic reticulum">Smooth endoplasmic reticulum</a></li> <li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a> (fluid that contains <a href="/wiki/Organelle" title="Organelle">organelles</a>; with which, comprises <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>)</li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Cell_membrane" title="Cell membrane">Cell membrane</a></li></ol></div> </td></tr></tbody></table> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:0317_Cytoskeletal_Components.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/0317_Cytoskeletal_Components.jpg/300px-0317_Cytoskeletal_Components.jpg" decoding="async" width="300" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/0317_Cytoskeletal_Components.jpg/450px-0317_Cytoskeletal_Components.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/0317_Cytoskeletal_Components.jpg/600px-0317_Cytoskeletal_Components.jpg 2x" data-file-width="1109" data-file-height="554" /></a><figcaption>The cytoskeleton consists of (a) microtubules, (b) microfilaments, and (c) intermediate filaments.<sup id="cite_ref-Openstax_Anatomy_&_Physiology_attribution_1-0" class="reference"><a href="#cite_note-Openstax_Anatomy_&_Physiology_attribution-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The <b>cytoskeleton</b> is a complex, dynamic network of interlinking <a href="/wiki/Protein_filament" title="Protein filament">protein filaments</a> present in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> of all <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a>, including those of <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>.<sup id="cite_ref-Hardin_2-0" class="reference"><a href="#cite_note-Hardin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, it extends from the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a> to the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> and is composed of similar proteins in the various organisms. It is composed of three main components: <a href="/wiki/Microfilaments" class="mw-redirect" title="Microfilaments">microfilaments</a>, <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a>, and <a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubules</a>, and these are all capable of rapid growth and or disassembly depending on the cell's requirements.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>A multitude of functions can be performed by the cytoskeleton. Its primary function is to give the cell its shape and mechanical resistance to deformation, and through association with extracellular <a href="/wiki/Connective_tissue" title="Connective tissue">connective tissue</a> and other cells it stabilizes entire tissues.<sup id="cite_ref-Alberts_4-0" class="reference"><a href="#cite_note-Alberts-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Herrmann_5-0" class="reference"><a href="#cite_note-Herrmann-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The cytoskeleton can also contract, thereby deforming the cell and the cell's environment and allowing <a href="/wiki/Cellular_migration" class="mw-redirect" title="Cellular migration">cells to migrate</a>.<sup id="cite_ref-Fletcher_6-0" class="reference"><a href="#cite_note-Fletcher-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Moreover, it is involved in many <a href="/wiki/Cell_signaling" title="Cell signaling">cell signaling</a> pathways and in the uptake of extracellular material (<a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>),<sup id="cite_ref-Geli1998_7-0" class="reference"><a href="#cite_note-Geli1998-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> the segregation of <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> during <a href="/wiki/Cell_division" title="Cell division">cellular division</a>,<sup id="cite_ref-Alberts_4-1" class="reference"><a href="#cite_note-Alberts-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> the <a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinesis</a> stage of cell division,<sup id="cite_ref-Wickstead2011_8-0" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> as scaffolding to organize the contents of the cell in space<sup id="cite_ref-Fletcher_6-1" class="reference"><a href="#cite_note-Fletcher-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> and in <a href="/wiki/Intracellular_transport" title="Intracellular transport">intracellular transport</a> (for example, the movement of <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a> and <a href="/wiki/Organelle" title="Organelle">organelles</a> within the cell)<sup id="cite_ref-Alberts_4-2" class="reference"><a href="#cite_note-Alberts-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> and can be a template for the construction of a <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a>.<sup id="cite_ref-Alberts_4-3" class="reference"><a href="#cite_note-Alberts-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Furthermore, it can form specialized structures, such as <a href="/wiki/Flagellum" title="Flagellum">flagella</a>, <a href="/wiki/Cilium" title="Cilium">cilia</a>, <a href="/wiki/Lamellipodia" class="mw-redirect" title="Lamellipodia">lamellipodia</a> and <a href="/wiki/Podosomes" class="mw-redirect" title="Podosomes">podosomes</a>. The structure, function and dynamic behavior of the cytoskeleton can be very different, depending on organism and cell type.<sup id="cite_ref-Alberts_4-4" class="reference"><a href="#cite_note-Alberts-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wickstead2011_8-1" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Even within one cell, the cytoskeleton can change through association with other proteins and the previous history of the network.<sup id="cite_ref-Fletcher_6-2" class="reference"><a href="#cite_note-Fletcher-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>A large-scale example of an action performed by the cytoskeleton is <a href="/wiki/Muscle_contraction" title="Muscle contraction">muscle contraction</a>. This is carried out by groups of highly specialized cells working together. A main component in the cytoskeleton that helps show the true function of this muscle contraction is the <a href="/wiki/Microfilament" title="Microfilament">microfilament</a>. Microfilaments are composed of the most abundant cellular protein known as actin.<sup id="cite_ref-Cooper-2000_10-0" class="reference"><a href="#cite_note-Cooper-2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> During contraction of a <a href="/wiki/Muscle" title="Muscle">muscle</a>, within each muscle cell, <a href="/wiki/Myosin" title="Myosin">myosin</a> molecular motors collectively exert forces on parallel <a href="/wiki/Actin" title="Actin">actin</a> filaments. Muscle contraction starts from nerve impulses which then causes increased amounts of calcium to be released from the <a href="/wiki/Sarcoplasmic_reticulum" title="Sarcoplasmic reticulum">sarcoplasmic reticulum</a>. Increases in calcium in the cytosol allows muscle contraction to begin with the help of two proteins, <a href="/wiki/Tropomyosin" title="Tropomyosin">tropomyosin</a> and <a href="/wiki/Troponin" title="Troponin">troponin</a>.<sup id="cite_ref-Cooper-2000_10-1" class="reference"><a href="#cite_note-Cooper-2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Tropomyosin inhibits the interaction between actin and myosin, while troponin senses the increase in calcium and releases the inhibition.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> This action contracts the muscle cell, and through the synchronous process in many muscle cells, the entire muscle. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1903, <a href="/wiki/Nikolai_Koltsov" title="Nikolai Koltsov">Nikolai K. Koltsov</a> proposed that the shape of cells was determined by a network of tubules that he termed the cytoskeleton. The concept of a protein mosaic that dynamically coordinated cytoplasmic biochemistry was proposed by Rudolph Peters in 1929<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> while the term (<i>cytosquelette</i>, in French) was first introduced by French embryologist <a href="/wiki/Paul_Wintrebert" title="Paul Wintrebert">Paul Wintrebert</a> in 1931.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>When the cytoskeleton was first introduced, it was thought to be an uninteresting gel-like substance that helped organelles stay in place.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Much research took place to try to understand the purpose of the cytoskeleton and its components. </p><p>Initially, it was thought that the cytoskeleton was exclusive to eukaryotes but in 1992 it was discovered to be present in prokaryotes as well. This discovery came after the realization that bacteria possess proteins that are homologous to tubulin and actin; the main components of the eukaryotic cytoskeleton.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Eukaryotic_cytoskeleton">Eukaryotic cytoskeleton</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=2" title="Edit section: Eukaryotic cytoskeleton"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">Eukaryotic</a> cells contain three main kinds of cytoskeletal filaments: <a href="/wiki/Microfilaments" class="mw-redirect" title="Microfilaments">microfilaments</a>, <a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubules</a>, and <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a>. In <a href="/wiki/Neuron" title="Neuron">neurons</a> the intermediate filaments are known as <a href="/wiki/Neurofilament" title="Neurofilament">neurofilaments</a>.<sup id="cite_ref-Taran_16-0" class="reference"><a href="#cite_note-Taran-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Each type is formed by the <a href="/wiki/Polymerization" title="Polymerization">polymerization</a> of a distinct type of <a href="/wiki/Protein_subunit" title="Protein subunit">protein subunit</a> and has its own characteristic shape and <a href="/wiki/Intracellular" class="mw-redirect" title="Intracellular">intracellular</a> distribution. Microfilaments are <a href="/wiki/Polymer" title="Polymer">polymers</a> of the protein <a href="/wiki/Actin" title="Actin">actin</a> and are 7 nm in diameter. Microtubules are composed of <a href="/wiki/Tubulin" title="Tubulin">tubulin</a> and are 25 nm in diameter. Intermediate filaments are composed of various proteins, depending on the type of cell in which they are found; they are normally 8-12 nm in diameter.<sup id="cite_ref-Hardin_2-1" class="reference"><a href="#cite_note-Hardin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The cytoskeleton provides the cell with structure and shape, and by <a href="/wiki/Excluded_volume" title="Excluded volume">excluding</a> <a href="/wiki/Macromolecules" class="mw-redirect" title="Macromolecules">macromolecules</a> from some of the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>, it adds to the level of <a href="/wiki/Macromolecular_crowding" title="Macromolecular crowding">macromolecular crowding</a> in this compartment.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Cytoskeletal elements interact extensively and intimately with cellular membranes.<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> </p><p>Research into <a href="/wiki/Neurodegeneration" class="mw-redirect" title="Neurodegeneration">neurodegenerative disorders</a> such as <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a>, <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a>, <a href="/wiki/Huntington%27s_disease" title="Huntington's disease">Huntington's disease</a>, and <a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">amyotrophic lateral sclerosis</a> (ALS) indicate that the cytoskeleton is affected in these diseases.<sup id="cite_ref-PelucchiStringhi2020_19-0" class="reference"><a href="#cite_note-PelucchiStringhi2020-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Parkinson's disease is marked by the degradation of neurons, resulting in tremors, rigidity, and other non-motor symptoms. Research has shown that microtubule assembly and stability in the cytoskeleton is compromised causing the neurons to degrade over time.<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> In Alzheimer's disease, <a href="/wiki/Tau_protein" title="Tau protein">tau proteins</a> which stabilize microtubules malfunction in the progression of the illness causing pathology of the cytoskeleton.<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> Excess glutamine in the Huntington protein involved with linking vesicles onto the cytoskeleton is also proposed to be a factor in the development of Huntington's Disease.<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> Amyotrophic lateral sclerosis results in a loss of movement caused by the degradation of motor neurons, and also involves defects of the cytoskeleton.<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><p><a href="/wiki/Stuart_Hameroff" title="Stuart Hameroff">Stuart Hameroff</a> and <a href="/wiki/Roger_Penrose" title="Roger Penrose">Roger Penrose</a> suggest a role of microtubule vibrations in <a href="/wiki/Neurons" class="mw-redirect" title="Neurons">neurons</a> in the origin of <a href="/wiki/Consciousness" title="Consciousness">consciousness</a>.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Accessory proteins including <a href="/wiki/Motor_protein" title="Motor protein">motor proteins</a> regulate and link the filaments to other cell compounds and each other and are essential for controlled assembly of cytoskeletal filaments in particular locations.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>A number of small-molecule <a href="/wiki/Cytoskeletal_drugs" title="Cytoskeletal drugs">cytoskeletal drugs</a> have been discovered that interact with actin and microtubules. These compounds have proven useful in studying the cytoskeleton, and several have clinical applications. </p> <div class="mw-heading mw-heading3"><h3 id="Microfilaments">Microfilaments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=3" title="Edit section: Microfilaments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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">Main article: <a href="/wiki/Microfilament" title="Microfilament">Microfilament</a></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:492px;max-width:492px"><div class="trow"><div class="tsingle" style="width:244px;max-width:244px"><div class="thumbimage" style="height:181px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Microfilament_Structure.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Microfilament_Structure.svg/242px-Microfilament_Structure.svg.png" decoding="async" width="242" height="182" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Microfilament_Structure.svg/363px-Microfilament_Structure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/96/Microfilament_Structure.svg/484px-Microfilament_Structure.svg.png 2x" data-file-width="512" data-file-height="384" /></a></span></div><div class="thumbcaption">Structure of a <a href="/wiki/Microfilament" title="Microfilament">microfilament</a></div></div><div class="tsingle" style="width:244px;max-width:244px"><div class="thumbimage" style="height:181px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:MEF_microfilaments.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/MEF_microfilaments.jpg/242px-MEF_microfilaments.jpg" decoding="async" width="242" height="182" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/MEF_microfilaments.jpg/363px-MEF_microfilaments.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e9/MEF_microfilaments.jpg/484px-MEF_microfilaments.jpg 2x" data-file-width="600" data-file-height="450" /></a></span></div><div class="thumbcaption">Actin cytoskeleton of <a href="/wiki/Mus_musculus" class="mw-redirect" title="Mus musculus">mouse</a> <a href="/wiki/Embryo" title="Embryo">embryo</a> <a href="/wiki/Fibroblast" title="Fibroblast">fibroblasts</a>, stained with <a href="/wiki/Phalloidin" title="Phalloidin">phalloidin</a></div></div></div></div></div> <p>Microfilaments, also known as actin filaments, are composed of linear polymers of <a href="/wiki/Actin#G-Actin" title="Actin">G-actin</a> proteins, and generate force when the growing (plus) end of the filament pushes against a barrier, such as the cell membrane. They also act as tracks for the movement of <a href="/wiki/Myosin" title="Myosin">myosin</a> molecules that affix to the microfilament and "walk" along them. In general, the major component or protein of microfilaments are actin. The G-actin monomer combines to form a polymer which continues to form the microfilament (actin filament). These subunits then assemble into two chains that intertwine into what are called <a href="/wiki/Actin#F-Actin" title="Actin">F-actin</a> chains.<sup id="cite_ref-Cooper_2000_27-0" class="reference"><a href="#cite_note-Cooper_2000-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Myosin motoring along F-actin filaments generates contractile forces in so-called actomyosin fibers, both in muscle as well as most non-muscle cell types.<sup id="cite_ref-gunning_28-0" class="reference"><a href="#cite_note-gunning-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Actin structures are controlled by the <a href="/wiki/Rho_family" class="mw-redirect" title="Rho family">Rho family</a> of small GTP-binding proteins such as Rho itself for contractile acto-myosin filaments ("stress fibers"), Rac for lamellipodia and Cdc42 for filopodia. </p><p>Functions include: </p> <ul><li><a href="/wiki/Muscle_contraction" title="Muscle contraction">Muscle contraction</a></li> <li>Cell movement</li> <li>Intracellular transport/trafficking</li> <li>Maintenance of <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cell shape</li> <li><a href="/wiki/Cytokinesis" title="Cytokinesis">Cytokinesis</a></li> <li>Cytoplasmic streaming<sup id="cite_ref-Cooper_2000_27-1" class="reference"><a href="#cite_note-Cooper_2000-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Intermediate_filaments">Intermediate filaments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=4" title="Edit section: Intermediate filaments"><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/Intermediate_filament" title="Intermediate filament">Intermediate filament</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:492px;max-width:492px"><div class="trow"><div class="tsingle" style="width:320px;max-width:320px"><div class="thumbimage" style="height:166px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Intermediate_filaments.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Intermediate_filaments.svg/318px-Intermediate_filaments.svg.png" decoding="async" width="318" height="166" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Intermediate_filaments.svg/477px-Intermediate_filaments.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Intermediate_filaments.svg/636px-Intermediate_filaments.svg.png 2x" data-file-width="512" data-file-height="268" /></a></span></div><div class="thumbcaption">Structure of an <a href="/wiki/Intermediate_filament" title="Intermediate filament">intermediate filament</a></div></div><div class="tsingle" style="width:168px;max-width:168px"><div class="thumbimage" style="height:166px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:KeratinF9.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/KeratinF9.png/166px-KeratinF9.png" decoding="async" width="166" height="167" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/KeratinF9.png/249px-KeratinF9.png 1.5x, //upload.wikimedia.org/wikipedia/commons/8/8e/KeratinF9.png 2x" data-file-width="308" data-file-height="310" /></a></span></div><div class="thumbcaption">Microscopy of <a href="/wiki/Keratin" title="Keratin">keratin</a> filaments inside cells</div></div></div></div></div> <p>Intermediate filaments are a part of the cytoskeleton of many <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells. These filaments, averaging 10 nanometers in diameter, are more stable (strongly bound) than microfilaments, and heterogeneous constituents of the cytoskeleton. Like <a href="/wiki/Actin" title="Actin">actin</a> filaments, they function in the maintenance of cell-shape by bearing tension (<a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubules</a>, by contrast, resist compression but can also bear tension during <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> and during the positioning of the centrosome). Intermediate filaments organize the internal tridimensional structure of the cell, anchoring <a href="/wiki/Organelle" title="Organelle">organelles</a> and serving as structural components of the <a href="/wiki/Nuclear_lamina" title="Nuclear lamina">nuclear lamina</a>. They also participate in some cell-cell and cell-matrix junctions. <a href="/wiki/Nuclear_lamina" title="Nuclear lamina">Nuclear lamina</a> exist in all animals and all tissues. Some animals like the <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">fruit fly</a> do not have any cytoplasmic intermediate filaments. In those animals that express cytoplasmic intermediate filaments, these are tissue specific.<sup id="cite_ref-Herrmann_5-1" class="reference"><a href="#cite_note-Herrmann-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Keratin intermediate filaments in <a href="/wiki/Epithelial" class="mw-redirect" title="Epithelial">epithelial</a> cells provide protection for different mechanical stresses the skin may endure. They also provide protection for organs against metabolic, oxidative, and chemical stresses. Strengthening of epithelial cells with these intermediate filaments may prevent onset of <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, or cell death, by reducing the probability of stress.<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> </p><p>Intermediate filaments are most commonly known as the support system or "scaffolding" for the cell and nucleus while also playing a role in some cell functions. In combination with proteins and <a href="/wiki/Desmosome" title="Desmosome">desmosomes</a>, the intermediate filaments form cell-cell connections and anchor the cell-matrix junctions that are used in messaging between cells as well as vital functions of the cell. These connections allow the cell to communicate through the desmosome of multiple cells to adjust structures of the tissue based on signals from the cells environment. Mutations in the IF proteins have been shown to cause serious medical issues such as premature aging, desmin mutations compromising organs, <a href="/wiki/Alexander_disease" title="Alexander disease">Alexander Disease</a>, and <a href="/wiki/Muscular_dystrophy" title="Muscular dystrophy">muscular dystrophy</a>.<sup id="cite_ref-Herrmann_5-2" class="reference"><a href="#cite_note-Herrmann-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Different intermediate filaments are: </p> <ul><li>made of <a href="/wiki/Vimentin" title="Vimentin">vimentins</a>. Vimentin intermediate filaments are in general present in mesenchymal cells.</li> <li>made of <a href="/wiki/Keratin" title="Keratin">keratin</a>. Keratin is present in general in epithelial cells.</li> <li><a href="/wiki/Neurofilament" title="Neurofilament">neurofilaments</a> of neural cells.</li> <li>made of <a href="/wiki/Lamin" title="Lamin">lamin</a>, giving structural support to the nuclear envelope.</li> <li>made of <a href="/wiki/Desmin" title="Desmin">desmin</a>, play an important role in structural and mechanical support of muscle cells.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Microtubules">Microtubules</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=5" title="Edit section: Microtubules"><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/Microtubule" title="Microtubule">Microtubule</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:492px;max-width:492px"><div class="trow"><div class="tsingle" style="width:245px;max-width:245px"><div class="thumbimage" style="height:215px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Microtubule_Structure.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Microtubule_Structure.svg/243px-Microtubule_Structure.svg.png" decoding="async" width="243" height="215" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Microtubule_Structure.svg/365px-Microtubule_Structure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Microtubule_Structure.svg/486px-Microtubule_Structure.svg.png 2x" data-file-width="512" data-file-height="454" /></a></span></div><div class="thumbcaption">Structure of a <a href="/wiki/Microtubule" title="Microtubule">microtubule</a></div></div><div class="tsingle" style="width:243px;max-width:243px"><div class="thumbimage" style="height:215px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Btub.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Btub.jpg/241px-Btub.jpg" decoding="async" width="241" height="215" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Btub.jpg/362px-Btub.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Btub.jpg/482px-Btub.jpg 2x" data-file-width="779" data-file-height="695" /></a></span></div><div class="thumbcaption">Microtubules in a gel-fixated cell</div></div></div></div></div> <p>Microtubules are hollow cylinders about 23 nm in diameter (lumen diameter of approximately 15 nm), most commonly comprising 13 <a href="/wiki/Microtubule" title="Microtubule">protofilaments</a> that, in turn, are polymers of alpha and beta <a href="/wiki/Tubulin" title="Tubulin">tubulin</a>. They have a very dynamic behavior, binding <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> for polymerization. They are commonly organized by the <a href="/wiki/Centrosome" title="Centrosome">centrosome</a>. </p><p>In nine triplet sets (star-shaped), they form the <a href="/wiki/Centrioles" class="mw-redirect" title="Centrioles">centrioles</a>, and in nine doublets oriented about two additional microtubules (wheel-shaped), they form cilia and flagella. The latter formation is commonly referred to as a "9+2" arrangement, wherein each doublet is connected to another by the protein <a href="/wiki/Dynein" title="Dynein">dynein</a>. As both flagella and cilia are structural components of the cell, and are maintained by microtubules, they can be considered part of the cytoskeleton. There are two types of cilia: motile and non-motile cilia. Cilia are short and more numerous than flagella. The motile cilia have a rhythmic waving or beating motion compared to the non-motile cilia which receive sensory information for the cell; processing signals from the other cells or the fluids surrounding it. Additionally, the microtubules control the beating (movement) of the cilia and flagella.<sup id="cite_ref-auto_31-0" class="reference"><a href="#cite_note-auto-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Also, the dynein arms attached to the microtubules function as the molecular motors. The motion of the cilia and flagella is created by the microtubules sliding past one another, which requires ATP.<sup id="cite_ref-auto_31-1" class="reference"><a href="#cite_note-auto-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> They play key roles in: </p> <ul><li>intracellular transport (associated with dyneins and <a href="/wiki/Kinesin" title="Kinesin">kinesins</a>, they transport <a href="/wiki/Organelles" class="mw-redirect" title="Organelles">organelles</a> like <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> or <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a>).</li> <li><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Bronchiolar_area_cilia_cross-sections_2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Bronchiolar_area_cilia_cross-sections_2.jpg/220px-Bronchiolar_area_cilia_cross-sections_2.jpg" decoding="async" width="220" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Bronchiolar_area_cilia_cross-sections_2.jpg/330px-Bronchiolar_area_cilia_cross-sections_2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Bronchiolar_area_cilia_cross-sections_2.jpg/440px-Bronchiolar_area_cilia_cross-sections_2.jpg 2x" data-file-width="1560" data-file-height="1257" /></a><figcaption>Cross section diagram through the cilium, showing the “9 + 2” arrangement of microtubules</figcaption></figure>the <a href="/wiki/Axoneme" title="Axoneme">axoneme</a> of <a href="/wiki/Cilium" title="Cilium">cilia</a> and <a href="/wiki/Flagellum" title="Flagellum">flagella</a>.</li> <li>the <a href="/wiki/Mitotic_spindle" class="mw-redirect" title="Mitotic spindle">mitotic spindle</a>.</li> <li>synthesis of the cell wall in plants.</li></ul> <p>In addition to the roles described above, Stuart Hameroff and Roger Penrose have proposed that microtubules function in consciousness.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Comparison">Comparison</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=6" title="Edit section: Comparison"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table class="wikitable"> <tbody><tr> <th>Cytoskeleton <br />type<sup id="cite_ref-boron25Unless_33-0" class="reference"><a href="#cite_note-boron25Unless-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </th> <th>Diameter <br />(<a href="/wiki/Nanometre" title="Nanometre">nm</a>)<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </th> <th>Structure </th> <th>Subunit examples<sup id="cite_ref-boron25Unless_33-1" class="reference"><a href="#cite_note-boron25Unless-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </th></tr> <tr> <th><a href="/wiki/Microfilaments" class="mw-redirect" title="Microfilaments">Microfilaments</a> </th> <td>6 </td> <td><a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">Double helix</a> </td> <td><a href="/wiki/Actin" title="Actin">Actin</a> </td></tr> <tr> <th><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate <br />filaments</a> </th> <td>10 </td> <td>Two anti-parallel <a href="/wiki/Helix" title="Helix">helices</a>/dimers, forming tetramers </td> <td> <ul><li><a href="/wiki/Vimentin" title="Vimentin">Vimentin</a> (<a href="/wiki/Mesenchyme" title="Mesenchyme">mesenchyme</a>)</li> <li><a href="/wiki/Glial_fibrillary_acidic_protein" title="Glial fibrillary acidic protein">Glial fibrillary acidic protein</a> (<a href="/wiki/Glial_cell" class="mw-redirect" title="Glial cell">glial cells</a>)</li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofilament</a> proteins (neuronal processes)</li> <li><a href="/wiki/Keratin" title="Keratin">Keratins</a> (<a href="/wiki/Epithelial_cell" class="mw-redirect" title="Epithelial cell">epithelial cells</a>)</li> <li><a href="/wiki/Nuclear_lamins" class="mw-redirect" title="Nuclear lamins">Nuclear lamins</a></li></ul> </td></tr> <tr> <th><a href="/wiki/Microtubule" title="Microtubule">Microtubules</a> </th> <td>23 </td> <td><a href="/wiki/Protofilament" class="mw-redirect" title="Protofilament">Protofilaments</a>, in turn consisting of tubulin subunits in complex with <a href="/wiki/Stathmin_protein_domain" class="mw-redirect" title="Stathmin protein domain">stathmin</a><sup id="cite_ref-pmid17029844_35-0" class="reference"><a href="#cite_note-pmid17029844-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </td> <td><a href="/wiki/%CE%91-bubulin" class="mw-redirect" title="Α-bubulin">α-</a> and <a href="/wiki/%CE%92-tubulin" class="mw-redirect" title="Β-tubulin">β-Tubulin</a> </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Septins">Septins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=7" title="Edit section: Septins"><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/Septin" title="Septin">Septin</a></div> <p>Septins are a group of the highly conserved <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> binding proteins found in <a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a>. Different septins form <a href="/wiki/Protein_complex" title="Protein complex">protein complexes</a> with each other. These can assemble to filaments and rings. Therefore, septins can be considered part of the cytoskeleton.<sup id="cite_ref-Mostowy2012_36-0" class="reference"><a href="#cite_note-Mostowy2012-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The function of septins in cells include serving as a localized attachment site for other <a href="/wiki/Protein" title="Protein">proteins</a>, and preventing the <a href="/wiki/Diffusion" title="Diffusion">diffusion</a> of certain molecules from one cell compartment to another.<sup id="cite_ref-Mostowy2012_36-1" class="reference"><a href="#cite_note-Mostowy2012-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> In yeast cells, they build scaffolding to provide structural support during cell division and compartmentalize parts of the cell. Recent research in human cells suggests that septins build cages around bacterial pathogens, immobilizing the harmful microbes and preventing them from invading other cells.<sup id="cite_ref-Mascarelli_2011_37-0" class="reference"><a href="#cite_note-Mascarelli_2011-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Spectrin">Spectrin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=8" title="Edit section: Spectrin"><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/Spectrin" title="Spectrin">Spectrin</a></div> <p>Spectrin is a cytoskeletal <a href="/wiki/Protein" title="Protein">protein</a> that lines the intracellular side of the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> in eukaryotic cells. Spectrin forms pentagonal or hexagonal arrangements, forming a <a href="/wiki/Scaffolding" title="Scaffolding">scaffolding</a> and playing an important role in maintenance of <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> integrity and cytoskeletal structure.<sup id="cite_ref-Huh_38-0" class="reference"><a href="#cite_note-Huh-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Yeast_cytoskeleton">Yeast cytoskeleton</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=9" title="Edit section: Yeast cytoskeleton"><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/Yeast" title="Yeast">Yeast</a></div> <p>In budding <a href="/wiki/Yeast" title="Yeast">yeast</a> (an important <a href="/wiki/Model_organism" title="Model organism">model organism</a>), <a href="/wiki/Actin" title="Actin">actin</a> forms cortical patches, actin cables, and a cytokinetic ring and the cap. Cortical patches are discrete actin bodies on the membrane and are vital for <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>, especially the recycling of glucan synthase which is important for <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a> synthesis. Actin cables are bundles of <a href="/wiki/Actin_filaments" class="mw-redirect" title="Actin filaments">actin filaments</a> and are involved in the transport of <a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">vesicles</a> towards the cap (which contains a number of different proteins to polarize cell growth) and in the positioning of mitochondria. The <a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinetic</a> ring forms and constricts around the site of <a href="/wiki/Cell_division" title="Cell division">cell division</a>.<sup id="cite_ref-Pruyne2000_39-0" class="reference"><a href="#cite_note-Pruyne2000-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Prokaryotic_cytoskeleton">Prokaryotic cytoskeleton</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=10" title="Edit section: Prokaryotic cytoskeleton"><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/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a></div> <p>Prior to the work of Jones et al., 2001, the cell wall was believed to be the deciding factor for many bacterial cell shapes, including rods and spirals. When studied, many misshapen bacteria were found to have mutations linked to development of a <a href="/wiki/Cell_envelope" title="Cell envelope">cell envelope</a>.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The cytoskeleton was once thought to be a feature only of <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells, but <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologues</a> to all the major proteins of the eukaryotic cytoskeleton have been found in <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>.<sup id="cite_ref-Shih_41-0" class="reference"><a href="#cite_note-Shih-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Harold Erickson notes that before 1992, only eukaryotes were believed to have cytoskeleton components. However, research in the early '90s suggested that bacteria and archaea had homologues of actin and tubulin, and that these were the basis of eukaryotic microtubules and microfilaments.<sup id="cite_ref-Erickson-2017_42-0" class="reference"><a href="#cite_note-Erickson-2017-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Although the evolutionary relationships are so distant that they are not obvious from protein sequence comparisons alone, the similarity of their three-dimensional <a href="/wiki/Protein_structure" title="Protein structure">structures</a> and similar functions in maintaining cell shape and polarity provides strong evidence that the eukaryotic and prokaryotic cytoskeletons are truly homologous.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Three laboratories independently discovered that FtsZ, a protein already known as a key player in bacterial cytokinesis, had the "tubulin signature sequence" present in all α-, β-, and γ-tubulins.<sup id="cite_ref-Erickson-2017_42-1" class="reference"><a href="#cite_note-Erickson-2017-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> However, some structures in the bacterial cytoskeleton may not have been identified as of yet.<sup id="cite_ref-gunning_28-1" class="reference"><a href="#cite_note-gunning-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="FtsZ">FtsZ</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=11" title="Edit section: FtsZ"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/FtsZ" title="FtsZ">FtsZ</a> was the first protein of the prokaryotic cytoskeleton to be identified. Like tubulin, FtsZ forms filaments in the presence of <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">guanosine triphosphate</a> (GTP), but these filaments do not group into tubules. During <a href="/wiki/Cell_division" title="Cell division">cell division</a>, FtsZ is the first protein to move to the division site, and is essential for recruiting other proteins that synthesize the new <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a> between the dividing cells. </p> <div class="mw-heading mw-heading3"><h3 id="MreB_and_ParM">MreB and ParM</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=12" title="Edit section: MreB and ParM"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Prokaryotic actin-like proteins, such as <a href="/wiki/MreB" title="MreB">MreB</a>, are involved in the maintenance of cell shape. All non-spherical bacteria have <a href="/wiki/Gene" title="Gene">genes</a> encoding actin-like proteins, and these proteins form a helical network beneath the cell membrane that guides the proteins involved in cell wall <a href="/wiki/Biosynthesis" title="Biosynthesis">biosynthesis</a>.<sup id="cite_ref-mreb_sheets_45-0" class="reference"><a href="#cite_note-mreb_sheets-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>Some <a href="/wiki/Plasmid" title="Plasmid">plasmids</a> encode a separate system that involves an actin-like protein <a href="/wiki/ParM" title="ParM">ParM</a>. Filaments of ParM exhibit <a href="/wiki/Microtubule#Dynamic_instability" title="Microtubule">dynamic instability</a>, and may partition plasmid DNA into the dividing daughter cells by a mechanism <a href="/wiki/Analogy_(biology)" class="mw-redirect" title="Analogy (biology)">analogous</a> to that used by microtubules during eukaryotic <a href="/wiki/Mitosis" title="Mitosis">mitosis</a>.<sup id="cite_ref-gunning_28-2" class="reference"><a href="#cite_note-gunning-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Alp_46-0" class="reference"><a href="#cite_note-Alp-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Crescentin">Crescentin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=13" title="Edit section: Crescentin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The bacterium <i><a href="/wiki/Caulobacter_crescentus" title="Caulobacter crescentus">Caulobacter crescentus</a></i> contains a third protein, <a href="/wiki/Crescentin" title="Crescentin">crescentin</a>, that is related to the intermediate filaments of eukaryotic cells. Crescentin is also involved in maintaining cell shape, such as helical and <a href="/wiki/Vibrio" title="Vibrio">vibrioid</a> forms of bacteria, but the mechanism by which it does this is currently unclear.<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> Additionally, curvature could be described by the displacement of crescentic filaments, after the disruption of peptidoglycan synthesis.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="The_cytoskeleton_and_cell_mechanics">The cytoskeleton and cell mechanics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=14" title="Edit section: The cytoskeleton and cell mechanics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cytoskeleton is a highly anisotropic and dynamic network, constantly remodeling itself in response to the changing cellular microenvironment. The network influences cell mechanics and dynamics by differentially polymerizing and depolymerizing its constituent filaments (primarily actin and myosin, but microtubules and intermediate filaments also play a role).<sup id="cite_ref-Chen_3285–3292_49-0" class="reference"><a href="#cite_note-Chen_3285–3292-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> This generates forces, which play an important role in informing the cell of its microenvironment. Specifically, forces such as tension, stiffness, and shear forces have all been shown to influence cell fate, differentiation, migration, and motility.<sup id="cite_ref-Chen_3285–3292_49-1" class="reference"><a href="#cite_note-Chen_3285–3292-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> Through a process called “mechanotransduction,” the cell remodels its cytoskeleton to sense and respond to these forces. </p><p><a href="/wiki/Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a> relies heavily on <a href="/wiki/Focal_adhesions" class="mw-redirect" title="Focal adhesions">focal adhesions</a>, which essentially connect the intracellular cytoskeleton with the <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> (ECM). Through focal adhesions, the cell is able to integrate extracellular forces into intracellular ones as the proteins present at focal adhesions undergo conformational changes to initiate signaling cascades. Proteins such as focal adhesion kinase (FAK) and Src have been shown to transduce force signals in response to cellular activities such as proliferation and differentiation, and are hypothesized to be key sensors in the mechanotransduction pathway.<sup id="cite_ref-Mechanisms_of_Mechanotransduction_50-0" class="reference"><a href="#cite_note-Mechanisms_of_Mechanotransduction-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> As a result of mechanotransduction, the cytoskeleton changes its composition and/or orientation to accommodate the force stimulus and ensure the cell responds accordingly. </p><p>The cytoskeleton changes the mechanics of the cell in response to detected forces. For example, increasing tension within the plasma membrane makes it more likely that ion channels will open, which increases ion conductance and makes cellular change ion influx or efflux much more likely.<sup id="cite_ref-Mechanisms_of_Mechanotransduction_50-1" class="reference"><a href="#cite_note-Mechanisms_of_Mechanotransduction-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Moreover, the mechanical properties of cells determine how far and where, directionally, a force will propagate throughout the cell and how it will change cell dynamics.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> A membrane protein that is not closely coupled to the cytoskeleton, for instance, will not produce a significant effect on the cortical actin network if it is subjected to a specifically directed force. However, membrane proteins that are more closely associated with the cytoskeleton will induce a more significant response.<sup id="cite_ref-Mechanisms_of_Mechanotransduction_50-2" class="reference"><a href="#cite_note-Mechanisms_of_Mechanotransduction-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> In this way, the anisotropy of the cytoskeleton serves to more keenly direct cell responses to intra or extracellular signals. </p> <div class="mw-heading mw-heading2"><h2 id="Long-range_order">Long-range order</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=15" title="Edit section: Long-range order"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The specific pathways and mechanisms by which the cytoskeleton senses and responds to forces are still under investigation. However, the <a href="/wiki/Long-range_order" class="mw-redirect" title="Long-range order">long-range order</a> generated by the cytoskeleton is known to contribute to mechanotransduction.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Cells, which are around 10–50 μm in diameter, are several thousand times larger than the molecules found within the cytoplasm that are essential to coordinate cellular activities. Because cells are so large in comparison to essential biomolecules, it is difficult, in the absence of an organizing network, for different parts of the cytoplasm to communicate.<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> Moreover, biomolecules must polymerize to lengths comparable to the length of the cell, but resulting polymers can be highly disorganized and unable to effectively transmit signals from one part of the cytoplasm to another. Thus, it is necessary to have the cytoskeleton to organize the polymers and ensure that they can effectively communicate across the entirety of the cell. </p> <div class="mw-heading mw-heading2"><h2 id="Common_features_and_differences_between_prokaryotes_and_eukaryotes">Common features and differences between prokaryotes and eukaryotes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=16" title="Edit section: Common features and differences between prokaryotes and eukaryotes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By definition, the cytoskeleton is composed of proteins that can form longitudinal arrays (fibres) in all organisms. These filament forming proteins have been classified into 4 classes. <a href="/wiki/Tubulin" title="Tubulin">Tubulin</a>-like, <a href="/wiki/Actin" title="Actin">actin</a>-like, Walker A cytoskeletal ATPases (WACA-proteins), and <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a>.<sup id="cite_ref-Wickstead2011_8-2" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-gunning_28-3" class="reference"><a href="#cite_note-gunning-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Tubulin-like proteins are <a href="/wiki/Tubulin" title="Tubulin">tubulin</a> in eukaryotes and <a href="/wiki/FtsZ" title="FtsZ">FtsZ</a>, TubZ, RepX in prokaryotes. Actin-like proteins are <a href="/wiki/Actin" title="Actin">actin</a> in eukaryotes and <a href="/wiki/MreB" title="MreB">MreB</a>, <a href="/wiki/FtsA" title="FtsA">FtsA</a> in prokaryotes. An example of a WACA-proteins, which are mostly found in prokaryotes, is <a href="/wiki/MinD" title="MinD">MinD</a>. Examples for intermediate filaments, which have almost exclusively been found in animals (i.e. eukaryotes) are the <a href="/wiki/Lamin" title="Lamin">lamins</a>, <a href="/wiki/Keratin" title="Keratin">keratins</a>, <a href="/wiki/Vimentin" title="Vimentin">vimentin</a>, <a href="/wiki/Neurofilament" title="Neurofilament">neurofilaments</a>, and <a href="/wiki/Desmin" title="Desmin">desmin</a>.<sup id="cite_ref-Wickstead2011_8-3" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Although tubulin-like proteins share some <a href="/wiki/Amino_acid_sequence" class="mw-redirect" title="Amino acid sequence">amino acid sequence</a> similarity, their equivalence in protein-fold and the similarity in the <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> binding site is more striking. The same holds true for the actin-like proteins and their structure and <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> binding domain.<sup id="cite_ref-Wickstead2011_8-4" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-gunning_28-4" class="reference"><a href="#cite_note-gunning-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Cytoskeletal proteins are usually correlated with cell shape, DNA segregation and cell division in prokaryotes and eukaryotes. Which proteins fulfill which task is very different. For example, DNA segregation in all eukaryotes happens through use of tubulin, but in prokaryotes either WACA proteins, actin-like or tubulin-like proteins can be used. Cell division is mediated in eukaryotes by actin, but in prokaryotes usually by tubulin-like (often FtsZ-ring) proteins and sometimes (<a href="/wiki/Thermoproteota" title="Thermoproteota">Thermoproteota</a>) <a href="/wiki/ESCRT#ESCRT-III" title="ESCRT">ESCRT-III</a>, which in eukaryotes still has a role in the last step of division.<sup id="cite_ref-Wickstead2011_8-5" class="reference"><a href="#cite_note-Wickstead2011-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Cytoplasmic_streaming">Cytoplasmic streaming</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=17" title="Edit section: Cytoplasmic streaming"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/220px--Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="124" data-durationhint="21" data-mwtitle="Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="854" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.720p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="720p.vp9.webm" data-width="1280" data-height="720" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.1080p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="1080p.vp9.webm" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm" type="video/webm; codecs="vp8"" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="256" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="426" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/9/95/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm/Movement_of_organelles_in_Tradescantia_stamen_hair_cells.webm.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="640" data-height="360" /></video></span><figcaption>Movement of organelles in <i><a href="/wiki/Tradescantia" title="Tradescantia">Tradescantia</a></i> stamen hair cells</figcaption></figure> <p><a href="/wiki/Cytoplasmic_streaming" title="Cytoplasmic streaming">Cytoplasmic streaming</a>, also known as cyclosis, is the active movement of a cell's contents along the components of the cytoskeleton. While mainly seen in plants, all cell types use this process for transportation of waste, nutrients, and organelles to other parts of the cell. <sup id="cite_ref-Woodhouse-2013_54-0" class="reference"><a href="#cite_note-Woodhouse-2013-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Plant and algae cells are generally larger than many other cells; so cytoplasmic streaming is important in these types of cells. This is because the cell's extra volume requires cytoplasmic streaming in order to move organelles throughout the entire cell.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Organelles move along <a href="/wiki/Microfilament" title="Microfilament">microfilaments</a> in the cytoskeleton driven by <a href="/wiki/Myosin" title="Myosin">myosin</a> motors binding and pushing along <a href="/wiki/Actin" title="Actin">actin</a> filament bundles.<sup id="cite_ref-Woodhouse-2013_54-1" class="reference"><a href="#cite_note-Woodhouse-2013-54"><span class="cite-bracket">[</span>54<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=Cytoskeleton&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span 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face of a cell membrane</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=Cytoskeleton&action=edit&section=19" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Openstax_Anatomy_&_Physiology_attribution-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Openstax_Anatomy_&_Physiology_attribution_1-0">^</a></b></span> <span class="reference-text"><i><span typeof="mw:File"><a href="//creativecommons.org/licenses/by/4.0/" title="creativecommons:by/4.0/"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Creative_Commons_by_small.svg/80px-Creative_Commons_by_small.svg.png" decoding="async" width="80" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Creative_Commons_by_small.svg/120px-Creative_Commons_by_small.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/65/Creative_Commons_by_small.svg/160px-Creative_Commons_by_small.svg.png 2x" data-file-width="80" data-file-height="15" /></a></span> This article incorporates <a rel="nofollow" class="external text" href="https://openstax.org/books/anatomy-and-physiology/pages/3-2-the-cytoplasm-and-cellular-organelles">text</a> available under the <a href="//creativecommons.org/licenses/by/4.0/" class="extiw" title="creativecommons:by/4.0/">CC BY 4.0</a> license.</i> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBettsDesaixJohnsonJohnson2023" class="citation book cs1">Betts, J Gordon; Desaix, Peter; Johnson, Eddie; Johnson, Jody E; Korol, Oksana; Kruse, Dean; Poe, Brandon; Wise, James; Womble, Mark D; Young, Kelly A (June 8, 2023). <i>Anatomy & Physiology</i>. Houston: OpenStax CNX. 3.2 The cytoplasm and cellular organelles. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-947172-04-3" title="Special:BookSources/978-1-947172-04-3"><bdi>978-1-947172-04-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Anatomy+%26+Physiology&rft.place=Houston&rft.pages=3.2+The+cytoplasm+and+cellular+organelles&rft.pub=OpenStax+CNX&rft.date=2023-06-08&rft.isbn=978-1-947172-04-3&rft.aulast=Betts&rft.aufirst=J+Gordon&rft.au=Desaix%2C+Peter&rft.au=Johnson%2C+Eddie&rft.au=Johnson%2C+Jody+E&rft.au=Korol%2C+Oksana&rft.au=Kruse%2C+Dean&rft.au=Poe%2C+Brandon&rft.au=Wise%2C+James&rft.au=Womble%2C+Mark+D&rft.au=Young%2C+Kelly+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACytoskeleton" class="Z3988"></span></span> </li> <li id="cite_note-Hardin-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Hardin_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Hardin_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHardinBertoniKleinsmith2015" class="citation book cs1">Hardin J, Bertoni G, Kleinsmith LJ (2015). <i>Becker's World of the Cell</i> (8th ed.). New York: Pearson. pp. 422–446. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978013399939-6" title="Special:BookSources/978013399939-6"><bdi>978013399939-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Becker%27s+World+of+the+Cell&rft.place=New+York&rft.pages=422-446&rft.edition=8th&rft.pub=Pearson&rft.date=2015&rft.isbn=978013399939-6&rft.aulast=Hardin&rft.aufirst=Jeff&rft.au=Bertoni%2C+Gregory&rft.au=Kleinsmith%2C+Lewis+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACytoskeleton" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcKinleyDean_O'LoughlinPennefather-O'BrienHarris2015" class="citation book cs1">McKinley, Michael; Dean O'Loughlin, Valerie; Pennefather-O'Brien, Elizabeth; Harris, Ronald (2015). <i>Human Anatomy</i> (4th ed.). New York: McGraw Hill Education. p. 29. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-07-352573-0" title="Special:BookSources/978-0-07-352573-0"><bdi>978-0-07-352573-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Human+Anatomy&rft.place=New+York&rft.pages=29&rft.edition=4th&rft.pub=McGraw+Hill+Education&rft.date=2015&rft.isbn=978-0-07-352573-0&rft.aulast=McKinley&rft.aufirst=Michael&rft.au=Dean+O%27Loughlin%2C+Valerie&rft.au=Pennefather-O%27Brien%2C+Elizabeth&rft.au=Harris%2C+Ronald&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACytoskeleton" class="Z3988"></span></span> </li> <li id="cite_note-Alberts-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Alberts_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Alberts_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Alberts_4-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Alberts_4-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Alberts_4-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlberts2008" class="citation book cs1">Alberts B, et al. (2008). <i>Molecular Biology of the Cell</i> (5th ed.). 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href="https://ui.adsabs.harvard.edu/abs/2013PNAS..11014132W">2013PNAS..11014132W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.1302736110">10.1073/pnas.1302736110</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3761564">3761564</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23940314">23940314</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Cytoplasmic+streaming+in+plant+cells+emerges+naturally+by+microfilament+self-organization&rft.volume=110&rft.issue=35&rft.pages=14132-7&rft.date=2013-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3761564%23id-name%3DPMC&rft_id=info%3Abibcode%2F2013PNAS..11014132W&rft_id=info%3Aarxiv%2F1308.6422&rft_id=info%3Apmid%2F23940314&rft_id=info%3Adoi%2F10.1073%2Fpnas.1302736110&rft.aulast=Woodhouse&rft.aufirst=FG&rft.au=Goldstein%2C+RE&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3761564&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACytoskeleton" class="Z3988"></span></span> </li> <li id="cite_note-55"><span class="mw-cite-backlink"><b><a href="#cite_ref-55">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGoldsteinvan_de_Meent2015" class="citation journal cs1">Goldstein RE, van de Meent JW (August 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4590424">"A physical perspective on cytoplasmic streaming"</a>. <i>Interface Focus</i>. <b>5</b> (4): 20150030. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsfs.2015.0030">10.1098/rsfs.2015.0030</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4590424">4590424</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26464789">26464789</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Interface+Focus&rft.atitle=A+physical+perspective+on+cytoplasmic+streaming&rft.volume=5&rft.issue=4&rft.pages=20150030&rft.date=2015-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4590424%23id-name%3DPMC&rft_id=info%3Apmid%2F26464789&rft_id=info%3Adoi%2F10.1098%2Frsfs.2015.0030&rft.aulast=Goldstein&rft.aufirst=RE&rft.au=van+de+Meent%2C+JW&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4590424&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACytoskeleton" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cytoskeleton&action=edit&section=20" title="Edit section: External links"><span>edit</span></a><span 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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 class="mw-selflink selflink">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 href="/wiki/Cilium" title="Cilium">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_the_cytoskeleton" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed 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:Cytoskeletal_proteins" title="Template:Cytoskeletal proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cytoskeletal_proteins" title="Template talk:Cytoskeletal proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cytoskeletal_proteins" title="Special:EditPage/Template:Cytoskeletal proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Proteins_of_the_cytoskeleton" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein" title="Protein">Proteins</a> of the <a class="mw-selflink selflink">cytoskeleton</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Human</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microfilament" title="Microfilament">Microfilaments</a><br />and <a href="/wiki/Actin-binding_protein" title="Actin-binding protein">ABPs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Myofilament" title="Myofilament">Myofilament</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Actin" title="Actin">Actins</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/Actin,_alpha_1" class="mw-redirect" title="Actin, alpha 1">A1</a></li> <li><a href="/wiki/ACTA2" title="ACTA2">A2</a></li> <li><a href="/wiki/Beta-actin" title="Beta-actin">B</a></li> <li><a href="/wiki/ACTC1" title="ACTC1">C1</a></li> <li><a href="/wiki/ACTG1" class="mw-redirect" title="ACTG1">G1</a></li> <li><a href="/wiki/ACTG2" title="ACTG2">G2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Myosin" title="Myosin">Myosins</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>I <ul><li><a href="/wiki/MYO1A" class="mw-redirect" title="MYO1A">MYO1A</a></li> <li><a href="/wiki/MYO1B" title="MYO1B">MYO1B</a></li> <li><a href="/wiki/MYO1C" title="MYO1C">MYO1C</a></li> <li><a href="/w/index.php?title=MYO1D&action=edit&redlink=1" class="new" title="MYO1D (page does not exist)">MYO1D</a></li> <li><a href="/wiki/MYO1E" title="MYO1E">MYO1E</a></li> <li><a href="/wiki/MYO1F" title="MYO1F">MYO1F</a></li> <li><a href="/wiki/MYO1G" title="MYO1G">MYO1G</a></li> <li><a href="/w/index.php?title=MYO1H&action=edit&redlink=1" class="new" title="MYO1H (page does not exist)">MYO1H</a>)</li></ul></li> <li>II <ul><li><a href="/wiki/MYH1" title="MYH1">MYH1</a></li> <li><a href="/wiki/MYH2" class="mw-redirect" title="MYH2">MYH2</a></li> <li><a href="/wiki/MYH3" title="MYH3">MYH3</a></li> <li><a href="/wiki/MYH4" title="MYH4">MYH4</a></li> <li><a href="/wiki/MYH6" title="MYH6">MYH6</a></li> <li><a href="/wiki/MYH7" title="MYH7">MYH7</a></li> <li><a href="/wiki/MYH7B" title="MYH7B">MYH7B</a></li> <li><a href="/wiki/MYH8" title="MYH8">MYH8</a></li> <li><a href="/wiki/MYH9" title="MYH9">MYH9</a></li> <li><a href="/wiki/MYH10" title="MYH10">MYH10</a></li> <li><a href="/wiki/MYH11" class="mw-redirect" title="MYH11">MYH11</a></li> <li><a href="/wiki/MYH13" title="MYH13">MYH13</a></li> <li><a href="/wiki/MYH14" title="MYH14">MYH14</a></li> <li><a href="/wiki/MYH15" title="MYH15">MYH15</a></li> <li><a href="/wiki/MYH16" class="mw-redirect" title="MYH16">MYH16</a></li></ul></li> <li>III <ul><li><a href="/wiki/MYO3A" class="mw-redirect" title="MYO3A">MYO3A</a></li> <li><a href="/w/index.php?title=MYO3B&action=edit&redlink=1" class="new" title="MYO3B (page does not exist)">MYO3B</a></li></ul></li> <li>V <ul><li><a href="/wiki/MYO5A" class="mw-redirect" title="MYO5A">MYO5A</a></li> <li><a href="/wiki/MYO5B" title="MYO5B">MYO5B</a></li> <li><a href="/w/index.php?title=MYO5C&action=edit&redlink=1" class="new" title="MYO5C (page does not exist)">MYO5C</a></li></ul></li> <li>VI <ul><li><a href="/wiki/MYO6" class="mw-redirect" title="MYO6">MYO6</a></li></ul></li> <li>VII <ul><li><a href="/wiki/MYO7A" title="MYO7A">MYO7A</a></li> <li><a href="/w/index.php?title=MYO7B&action=edit&redlink=1" class="new" title="MYO7B (page does not exist)">MYO7B</a></li></ul></li> <li>IX <ul><li><a href="/w/index.php?title=MYO9A&action=edit&redlink=1" class="new" title="MYO9A (page does not exist)">MYO9A</a></li> <li><a href="/wiki/MYO9B" title="MYO9B">MYO9B</a></li></ul></li> <li>X <ul><li><a href="/wiki/MYO10" title="MYO10">MYO10</a></li></ul></li> <li>XV <ul><li><a href="/wiki/MYO15A" title="MYO15A">MYO15A</a></li></ul></li> <li>XVIII <ul><li><a href="/wiki/MYO18A" title="MYO18A">MYO18A</a></li> <li><a href="/wiki/MYO18B" title="MYO18B">MYO18B</a></li></ul></li> <li><a href="/wiki/Myosin_light_chain" title="Myosin light chain">LC</a> <ul><li><a href="/wiki/MYL1" title="MYL1">MYL1</a></li> <li><a href="/wiki/MYL2" title="MYL2">MYL2</a></li> <li><a href="/wiki/MYL3" title="MYL3">MYL3</a></li> <li><a href="/wiki/MYL4" title="MYL4">MYL4</a></li> <li><a href="/w/index.php?title=MYL5&action=edit&redlink=1" class="new" title="MYL5 (page does not exist)">MYL5</a></li> <li><a href="/wiki/MYL6" title="MYL6">MYL6</a></li> <li><a href="/wiki/MYL6B" title="MYL6B">MYL6B</a></li> <li><a href="/wiki/MYL7" title="MYL7">MYL7</a></li> <li><a href="/wiki/MYL9" title="MYL9">MYL9</a></li> <li><a href="/wiki/MYLIP" title="MYLIP">MYLIP</a></li> <li><a href="/wiki/MYLK" title="MYLK">MYLK</a></li> <li><a href="/wiki/MYLK2" title="MYLK2">MYLK2</a></li> <li><a href="/w/index.php?title=MYLL1&action=edit&redlink=1" class="new" title="MYLL1 (page does not exist)">MYLL1</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Tropomodulin" title="Tropomodulin">Tropomodulin</a> <ul><li><a href="/wiki/TMOD1" class="mw-redirect" title="TMOD1">1</a></li> <li><a href="/wiki/TMOD2" class="mw-redirect" title="TMOD2">2</a></li> <li><a href="/wiki/TMOD3" class="mw-redirect" title="TMOD3">3</a></li> <li><a href="/wiki/TMOD4" class="mw-redirect" title="TMOD4">4</a></li></ul></li> <li><a href="/wiki/Troponin" title="Troponin">Troponin</a> <ul><li><a href="/wiki/Troponin_T" title="Troponin T">T</a> <a href="/wiki/TNNT1" title="TNNT1">1</a> <a href="/wiki/TNNT2" title="TNNT2">2</a> <a href="/wiki/TNNT3" title="TNNT3">3</a></li> <li><a href="/wiki/Troponin_C" title="Troponin C">C</a> <a href="/wiki/Troponin_C_type_1" title="Troponin C type 1">1</a> <a href="/wiki/TNNC2" class="mw-redirect" title="TNNC2">2</a></li> <li><a href="/wiki/Troponin_I" title="Troponin I">I</a> <a href="/wiki/TNNI1" title="TNNI1">1</a> <a href="/wiki/TNNI2" title="TNNI2">2</a> <a href="/wiki/TNNI3" title="TNNI3">3</a></li></ul></li> <li><a href="/wiki/Tropomyosin" title="Tropomyosin">Tropomyosin</a> <ul><li><a href="/wiki/TPM1" title="TPM1">1</a></li> <li><a href="/wiki/TPM2" title="TPM2">2</a></li> <li><a href="/wiki/Tropomyosin_3" title="Tropomyosin 3">3</a></li> <li><a href="/wiki/TPM4" title="TPM4">4</a></li></ul></li></ul> <ul><li><a href="/wiki/Actinin" title="Actinin">Actinin</a> <ul><li><a href="/wiki/Actinin,_alpha_1" class="mw-redirect" title="Actinin, alpha 1">1</a></li> <li><a href="/wiki/Actinin,_alpha_2" class="mw-redirect" title="Actinin, alpha 2">2</a></li> <li><a href="/wiki/ACTN3" class="mw-redirect" title="ACTN3">3</a></li> <li><a href="/wiki/Actinin_alpha_4" class="mw-redirect" title="Actinin alpha 4">4</a></li></ul></li> <li><a href="/wiki/Arp2/3_complex" title="Arp2/3 complex">Arp2/3 complex</a></li> <li><i><a href="/wiki/Actin_depolymerizing_factor" class="mw-redirect" title="Actin depolymerizing factor">actin depolymerizing factors</a></i> <ul><li><a href="/wiki/Cofilin" class="mw-redirect" title="Cofilin">Cofilin</a> <ul><li><a href="/wiki/Cofilin_1" title="Cofilin 1">1</a></li> <li><a href="/wiki/CFL2_(gene)" class="mw-redirect" title="CFL2 (gene)">2</a></li></ul></li> <li><a href="/wiki/Destrin" title="Destrin">Destrin</a></li></ul></li> <li><a href="/wiki/Gelsolin" title="Gelsolin">Gelsolin</a></li> <li><a href="/wiki/Profilin" title="Profilin">Profilin</a> <ul><li><a href="/wiki/Profilin_1" title="Profilin 1">1</a></li> <li><a href="/wiki/PFN2" title="PFN2">2</a></li></ul></li> <li><a href="/wiki/Titin" title="Titin">Titin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Wiskott%E2%80%93Aldrich_syndrome_protein" title="Wiskott–Aldrich syndrome protein">Wiskott–Aldrich syndrome protein</a></li> <li><a href="/wiki/Fibrillin" title="Fibrillin">Fibrillin</a></li> <li><a href="/wiki/Filamin" title="Filamin">Filamin</a> <ul><li><a href="/wiki/FLNA" title="FLNA">FLNA</a></li> <li><a href="/wiki/FLNB" title="FLNB">FLNB</a></li> <li><a href="/wiki/FLNC_(gene)" title="FLNC (gene)">FLNC</a></li></ul></li> <li><a href="/wiki/Espin_(protein)" title="Espin (protein)">Espin</a></li> <li><a href="/wiki/TRIOBP" title="TRIOBP">TRIOBP</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/Intermediate_filament" title="Intermediate filament">Intermediate <br />filaments</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%">Type 1/2<br />(<a href="/wiki/Keratin" title="Keratin">Keratin</a>,<br /><a href="/wiki/Cytokeratin" title="Cytokeratin">Cytokeratin</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Epithelial_keratin" class="mw-redirect" title="Epithelial keratin">Epithelial keratins</a><br />(soft alpha-keratins)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Type_I_keratin" title="Type I keratin">type I</a>/<a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/Keratin_9" title="Keratin 9">9</a></li> <li><a href="/wiki/Keratin_10" title="Keratin 10">10</a></li> <li><a href="/wiki/Keratin_12" title="Keratin 12">12</a></li> <li><a href="/wiki/Keratin_13" title="Keratin 13">13</a></li> <li><a href="/wiki/Keratin_14" title="Keratin 14">14</a></li> <li><a href="/wiki/Keratin_15" title="Keratin 15">15</a></li> <li><a href="/wiki/Keratin_16" title="Keratin 16">16</a></li> <li><a href="/wiki/Keratin_17" title="Keratin 17">17</a></li> <li><a href="/wiki/Keratin_19" title="Keratin 19">19</a></li> <li><a href="/wiki/Keratin_20" title="Keratin 20">20</a></li></ul></li> <li><i><a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/Keratin_18" title="Keratin 18">18</a></li></ul></li> <li><i>none</i> <ul><li><a href="/wiki/Keratin_21" title="Keratin 21">21</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Type_II_keratin" title="Type II keratin">type II</a>/<a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/Keratin_1" title="Keratin 1">1</a></li> <li><a href="/wiki/Keratin_2A" title="Keratin 2A">2A</a></li> <li><a href="/wiki/Keratin_3" title="Keratin 3">3</a></li> <li><a href="/wiki/Keratin_4" title="Keratin 4">4</a></li> <li><a href="/wiki/Keratin_5" title="Keratin 5">5</a></li> <li><a href="/wiki/Keratin_6A" title="Keratin 6A">6A</a></li> <li><a href="/wiki/Keratin_6B" title="Keratin 6B">6B</a></li> <li><a href="/wiki/Keratin_6C" title="Keratin 6C">6C</a></li> <li><a href="/wiki/Keratin_7" title="Keratin 7">7</a></li> <li><a href="/wiki/Keratin_8" title="Keratin 8">8</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hair_keratin" title="Hair keratin">Hair keratins</a><br />(hard alpha-keratins)</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Type_I_hair_keratin" class="mw-redirect" title="Type I hair keratin">type I</a>/<a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/KRT31" title="KRT31">31</a></li> <li><a href="/wiki/KRT32" title="KRT32">32</a></li> <li><a href="/wiki/KRT33A" title="KRT33A">33A</a></li> <li><a href="/wiki/KRT33B" title="KRT33B">33B</a></li> <li><a href="/wiki/Keratin_34" title="Keratin 34">34</a></li> <li><a href="/wiki/KRT35" title="KRT35">35</a></li> <li><a href="/wiki/KRT36" title="KRT36">36</a></li> <li><a href="/wiki/KRT37" title="KRT37">37</a></li> <li><a href="/wiki/KRT38" title="KRT38">38</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Type_II_hair_keratin" class="mw-redirect" title="Type II hair keratin">type II</a>/<a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/KRT81" title="KRT81">81</a></li> <li><a href="/wiki/KRT82" title="KRT82">82</a></li> <li><a href="/wiki/KRT83" title="KRT83">83</a></li> <li><a href="/wiki/KRT84" title="KRT84">84</a></li> <li><a href="/wiki/KRT85" title="KRT85">85</a></li> <li><a href="/wiki/KRT86" title="KRT86">86</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ungrouped alpha</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/KRT23" title="KRT23">23</a></li> <li><a href="/wiki/KRT24" title="KRT24">24</a></li> <li><a href="/wiki/KRT25" title="KRT25">25</a></li> <li><a href="/wiki/KRT26" title="KRT26">26</a></li> <li><a href="/wiki/KRT27" title="KRT27">27</a></li> <li><a href="/wiki/KRT28" title="KRT28">28</a></li> <li><a href="/wiki/KRT39" title="KRT39">39</a></li> <li><a href="/wiki/KRT40" title="KRT40">40</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/KRT71" title="KRT71">71</a></li> <li><a href="/wiki/KRT72" title="KRT72">72</a></li> <li><a href="/wiki/KRT73" title="KRT73">73</a></li> <li><a href="/wiki/KRT74" title="KRT74">74</a></li> <li><a href="/wiki/KRT75" title="KRT75">75</a></li> <li><a href="/wiki/KRT76" title="KRT76">76</a></li> <li><a href="/wiki/KRT77" title="KRT77">77</a></li> <li><a href="/wiki/KRT78" title="KRT78">78</a></li> <li><a href="/wiki/KRT79" title="KRT79">79</a></li> <li><a href="/wiki/KRT80" title="KRT80">80</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Not alpha</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/Beta-keratin" title="Beta-keratin">Beta-keratin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 3</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/Desmin" title="Desmin">Desmin</a></li> <li><a href="/wiki/Glial_fibrillary_acidic_protein" title="Glial fibrillary acidic protein">GFAP</a></li> <li><a href="/wiki/Peripherin" title="Peripherin">Peripherin</a></li> <li><a href="/wiki/Vimentin" title="Vimentin">Vimentin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 4</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/Internexin" title="Internexin">Internexin</a></li> <li><a href="/wiki/Nestin_(protein)" title="Nestin (protein)">Nestin</a></li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofilament</a> <ul><li><a href="/wiki/NEFL" class="mw-redirect" title="NEFL">NEFL</a></li> <li><a href="/wiki/NEFM" title="NEFM">NEFM</a></li> <li><a href="/wiki/NEFH" title="NEFH">NEFH</a></li></ul></li> <li><a href="/wiki/Synemin" title="Synemin">Synemin</a></li> <li><a href="/wiki/Syncoilin" title="Syncoilin">Syncoilin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 5</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/Lamin" title="Lamin">Nuclear lamins</a>: <a href="/wiki/Lamin_A/C" class="mw-redirect" title="Lamin A/C">A/C</a></li> <li><a href="/wiki/Lamin_B1" title="Lamin B1">B1</a></li> <li><a href="/wiki/Lamin_B2" title="Lamin B2">B2</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/Microtubule" title="Microtubule">Microtubules</a><br />and <a href="/wiki/Microtubule-associated_protein" title="Microtubule-associated protein">MAPs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tubulin" title="Tubulin">Tubulins</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/TUBA1A" class="mw-redirect" title="TUBA1A">TUBA1A</a></li> <li><a href="/wiki/TUBA1B" title="TUBA1B">TUBA1B</a></li> <li><a href="/wiki/TUBA1C" title="TUBA1C">TUBA1C</a></li> <li><a href="/wiki/TUBA3C" class="mw-redirect" title="TUBA3C">TUBA3C</a></li> <li><a href="/w/index.php?title=TUBA3D&action=edit&redlink=1" class="new" title="TUBA3D (page does not exist)">TUBA3D</a></li> <li><a href="/w/index.php?title=TUBA3E&action=edit&redlink=1" class="new" title="TUBA3E (page does not exist)">TUBA3E</a></li> <li><a href="/wiki/TUBA4A" title="TUBA4A">TUBA4A</a></li> <li><a href="/wiki/TUBA8" title="TUBA8">TUBA8</a></li> <li><a href="/wiki/TUBB" title="TUBB">TUBB</a></li> <li><a href="/wiki/TUBB1" title="TUBB1">TUBB1</a></li> <li><a href="/wiki/TUBB2A" class="mw-redirect" title="TUBB2A">TUBB2A</a></li> <li><a href="/w/index.php?title=TUBB2B&action=edit&redlink=1" class="new" title="TUBB2B (page does not exist)">TUBB2B</a></li> <li><a href="/wiki/TUBB2C" class="mw-redirect" title="TUBB2C">TUBB2C</a></li> <li><a href="/wiki/TUBB3" class="mw-redirect" title="TUBB3">TUBB3</a></li> <li><a href="/wiki/TUBB4" class="mw-redirect" title="TUBB4">TUBB4</a></li> <li><a href="/w/index.php?title=TUBB4Q&action=edit&redlink=1" class="new" title="TUBB4Q (page does not exist)">TUBB4Q</a></li> <li><a href="/w/index.php?title=TUBB6&action=edit&redlink=1" class="new" title="TUBB6 (page does not exist)">TUBB6</a></li> <li><a href="/w/index.php?title=TUBB8&action=edit&redlink=1" class="new" title="TUBB8 (page does not exist)">TUBB8</a></li> <li><a href="/wiki/TUBG1" title="TUBG1">TUBG1</a></li> <li><a href="/wiki/TUBG2" title="TUBG2">TUBG2</a></li> <li><a href="/wiki/TUBGCP2" title="TUBGCP2">TUBGCP2</a></li> <li><a href="/wiki/TUBGCP3" title="TUBGCP3">TUBGCP3</a></li> <li><a href="/wiki/TUBGCP4" title="TUBGCP4">TUBGCP4</a></li> <li><a href="/wiki/TUBGCP5" title="TUBGCP5">TUBGCP5</a></li> <li><a href="/wiki/TUBGCP6" title="TUBGCP6">TUBGCP6</a></li> <li><a href="/wiki/TUBD1" title="TUBD1">TUBD1</a></li> <li><a href="/wiki/TUBE1" title="TUBE1">TUBE1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microtubule-associated_protein" title="Microtubule-associated protein">MAPs</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/MAPRE1" title="MAPRE1">EB1</a></li> <li><a href="/wiki/MAPRE2" title="MAPRE2">EB2</a></li> <li><a href="/wiki/MAPRE3" title="MAPRE3">EB3</a></li> <li><a href="/wiki/MAP1A" title="MAP1A">MAP1A</a></li> <li><a href="/wiki/MAP1B" title="MAP1B">MAP1B</a></li> <li><a href="/wiki/MAP2" class="mw-redirect" title="MAP2">MAP2</a></li> <li><a href="/wiki/MAP4" title="MAP4">MAP4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Kinesin" title="Kinesin">Kinesins</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/KIF1A" title="KIF1A">KIF1A</a></li> <li><a href="/wiki/KIF1B" title="KIF1B">KIF1B</a></li> <li><a href="/wiki/KIF2A" title="KIF2A">KIF2A</a></li> <li><a href="/wiki/KIF2C" title="KIF2C">KIF2C</a></li> <li><a href="/wiki/KIF3B" title="KIF3B">KIF3B</a></li> <li><a href="/wiki/KIF3C" title="KIF3C">KIF3C</a></li> <li><a href="/wiki/KIF4A" title="KIF4A">KIF4A</a></li> <li><a href="/wiki/KIF4B" title="KIF4B">KIF4B</a></li> <li><a href="/wiki/KIF5A" title="KIF5A">KIF5A</a></li> <li><a href="/wiki/KIF5B" title="KIF5B">KIF5B</a></li> <li><a href="/wiki/KIF5C" title="KIF5C">KIF5C</a></li> <li><a href="/wiki/KIF6" title="KIF6">KIF6</a></li> <li><a href="/wiki/KIF7" title="KIF7">KIF7</a></li> <li><a href="/wiki/KIF9" title="KIF9">KIF9</a></li> <li><a href="/wiki/KIF11" class="mw-redirect" title="KIF11">KIF11</a></li> <li><a href="/wiki/KIF12" title="KIF12">KIF12</a></li> <li><a href="/wiki/KIF13A" title="KIF13A">KIF13A</a></li> <li><a href="/wiki/KIF13B" title="KIF13B">KIF13B</a></li> <li><a href="/wiki/KIF14" title="KIF14">KIF14</a></li> <li><a href="/wiki/KIF15" title="KIF15">KIF15</a></li> <li><a href="/wiki/KIF16B" title="KIF16B">KIF16B</a></li> <li><a href="/wiki/KIF17" title="KIF17">KIF17</a></li> <li><a href="/wiki/KIF18A" title="KIF18A">KIF18A</a></li> <li><a href="/wiki/KIF18B" title="KIF18B">KIF18B</a></li> <li><a href="/wiki/KIF19" title="KIF19">KIF19</a></li> <li><a href="/wiki/KIF20A" title="KIF20A">KIF20A</a></li> <li><a href="/wiki/KIF20B" title="KIF20B">KIF20B</a></li> <li><a href="/wiki/KIF21A" title="KIF21A">KIF21A</a></li> <li><a href="/wiki/KIF21B" title="KIF21B">KIF21B</a></li> <li><a href="/wiki/KIF22" title="KIF22">KIF22</a></li> <li><a href="/wiki/KIF23" title="KIF23">KIF23</a></li> <li><a href="/wiki/KIF24" title="KIF24">KIF24</a></li> <li><a href="/wiki/KIF25" title="KIF25">KIF25</a></li> <li><a href="/wiki/KIF26A" title="KIF26A">KIF26A</a></li> <li><a href="/w/index.php?title=KIF26B&action=edit&redlink=1" class="new" title="KIF26B (page does not exist)">KIF26B</a></li> <li><a href="/wiki/KIF27" title="KIF27">KIF27</a></li> <li><a href="/wiki/KIFC1" title="KIFC1">KIFC1</a></li> <li><a href="/w/index.php?title=KIFC2&action=edit&redlink=1" class="new" title="KIFC2 (page does not exist)">KIFC2</a></li> <li><a href="/wiki/KIFC3" title="KIFC3">KIFC3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dynein" title="Dynein">Dyneins</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>axonemal: <a href="/wiki/DNAH1" title="DNAH1">DNAH1</a></li> <li><a href="/w/index.php?title=DNAH2&action=edit&redlink=1" class="new" title="DNAH2 (page does not exist)">DNAH2</a></li> <li><a href="/w/index.php?title=DNAH3&action=edit&redlink=1" class="new" title="DNAH3 (page does not exist)">DNAH3</a></li> <li><a href="/wiki/DNAH5" title="DNAH5">DNAH5</a></li> <li><a href="/w/index.php?title=DNAH6&action=edit&redlink=1" class="new" title="DNAH6 (page does not exist)">DNAH6</a></li> <li><a href="/wiki/DNAH7" title="DNAH7">DNAH7</a></li> <li><a href="/w/index.php?title=DNAH8&action=edit&redlink=1" class="new" title="DNAH8 (page does not exist)">DNAH8</a></li> <li><a href="/wiki/DNAH9" title="DNAH9">DNAH9</a></li> <li><a href="/w/index.php?title=DNAH10&action=edit&redlink=1" class="new" title="DNAH10 (page does not exist)">DNAH10</a></li> <li><a href="/wiki/DNAH11" title="DNAH11">DNAH11</a></li> <li><a href="/w/index.php?title=DNAH12&action=edit&redlink=1" class="new" title="DNAH12 (page does not exist)">DNAH12</a></li> <li><a href="/w/index.php?title=DNAH13&action=edit&redlink=1" class="new" title="DNAH13 (page does not exist)">DNAH13</a></li> <li><a href="/w/index.php?title=DNAH14&action=edit&redlink=1" class="new" title="DNAH14 (page does not exist)">DNAH14</a></li> <li><a href="/w/index.php?title=DNAH17&action=edit&redlink=1" class="new" title="DNAH17 (page does not exist)">DNAH17</a></li> <li><a href="/wiki/DNAI1" title="DNAI1">DNAI1</a></li> <li><a href="/wiki/DNAI2" title="DNAI2">DNAI2</a></li> <li><a href="/w/index.php?title=DNALI1&action=edit&redlink=1" class="new" title="DNALI1 (page does not exist)">DNALI1</a></li> <li><a href="/wiki/DNAL1" class="mw-redirect" title="DNAL1">DNAL1</a></li> <li><a href="/wiki/DNAL4" title="DNAL4">DNAL4</a></li></ul> <ul><li>cytoplasmic: <a href="/wiki/DYNC1H1" title="DYNC1H1">DYNC1H1</a></li> <li><a href="/wiki/DYNC2H1" title="DYNC2H1">DYNC2H1</a></li> <li><a href="/wiki/DYNC1I1" title="DYNC1I1">DYNC1I1</a></li> <li><a href="/wiki/DYNC1I2" title="DYNC1I2">DYNC1I2</a></li> <li><a href="/wiki/DYNC1LI1" title="DYNC1LI1">DYNC1LI1</a></li> <li><a href="/wiki/DYNC1LI2" title="DYNC1LI2">DYNC1LI2</a></li> <li><a href="/w/index.php?title=DYNC2LI1&action=edit&redlink=1" class="new" title="DYNC2LI1 (page does not exist)">DYNC2LI1</a></li> <li><a href="/wiki/DYNLL1" title="DYNLL1">DYNLL1</a></li> <li><a href="/wiki/DYNLL2" title="DYNLL2">DYNLL2</a></li> <li><a href="/wiki/DYNLRB1" title="DYNLRB1">DYNLRB1</a></li> <li><a href="/w/index.php?title=DYNLRB2&action=edit&redlink=1" class="new" title="DYNLRB2 (page does not exist)">DYNLRB2</a></li> <li><a href="/wiki/DYNLT1" title="DYNLT1">DYNLT1</a></li> <li><a href="/wiki/DYNLT3" title="DYNLT3">DYNLT3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organising proteins</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/CAMSAP1" title="CAMSAP1">CAMSAP1</a></li> <li><a href="/wiki/CAMSAP2" title="CAMSAP2">CAMSAP2</a></li> <li><a href="/wiki/CAMSAP3" title="CAMSAP3">CAMSAP3</a></li> <li><a href="/wiki/Centrin_1" title="Centrin 1">Centrin 1</a></li> <li><a href="/wiki/Centrin_2" title="Centrin 2">Centrin 2</a></li> <li><a href="/wiki/Centrin_3" title="Centrin 3">Centrin 3</a></li> <li><a href="/wiki/PCM1" title="PCM1">PCM1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Microtubule severing proteins</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/Katanin" title="Katanin">Katanin</a></li> <li><a href="/wiki/Spastin" title="Spastin">Spastin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Tau_protein" title="Tau protein">Tau protein</a></li> <li><a href="/wiki/Dynactin" title="Dynactin">Dynactin</a> <ul><li><a href="/wiki/DCTN1" title="DCTN1">DCTN1</a></li></ul></li> <li><a href="/wiki/Stathmin" title="Stathmin">Stathmin</a></li> <li><a href="/wiki/Tektin" title="Tektin">Tektin</a> <ul><li><a href="/wiki/TEKT1" class="mw-redirect" title="TEKT1">TEKT1</a></li> <li><a href="/w/index.php?title=TEKT2&action=edit&redlink=1" class="new" title="TEKT2 (page does not exist)">TEKT2</a></li> <li><a href="/w/index.php?title=TEKT3&action=edit&redlink=1" class="new" title="TEKT3 (page does not exist)">TEKT3</a></li> <li><a href="/w/index.php?title=TEKT4&action=edit&redlink=1" class="new" title="TEKT4 (page does not exist)">TEKT4</a></li> <li><a href="/w/index.php?title=TEKT5&action=edit&redlink=1" class="new" title="TEKT5 (page does not exist)">TEKT5</a></li></ul></li> <li><a href="/wiki/Dynamin" title="Dynamin">Dynamin</a> <ul><li><a href="/wiki/DNM1" title="DNM1">DNM1</a></li> <li><a href="/wiki/DNM2" title="DNM2">DNM2</a></li> <li><a href="/wiki/DNM3" title="DNM3">DNM3</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Catenin" title="Catenin">Catenins</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/%CE%91-Catenin" title="Α-Catenin">Alpha catenin</a></li> <li><a href="/wiki/Catenin_beta-1" title="Catenin beta-1">Beta catenin</a> <ul><li><a href="/wiki/Adenomatous_polyposis_coli" title="Adenomatous polyposis coli">APC</a></li></ul></li> <li><a href="/wiki/Plakoglobin" title="Plakoglobin">Plakoglobin (gamma catenin)</a></li> <li><a href="/wiki/%CE%94-Catenin" title="Δ-Catenin">Delta catenin</a></li> <li><a href="/wiki/Gigaxonin" title="Gigaxonin">GAN</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Membrane_protein" title="Membrane protein">Membrane</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/Dystrophin" title="Dystrophin">Dystrophin</a> <ul><li><a href="/wiki/Dystroglycan" title="Dystroglycan">Dystroglycan</a></li></ul></li> <li><a href="/wiki/Utrophin" title="Utrophin">Utrophin</a></li> <li><a href="/wiki/Ankyrin" title="Ankyrin">Ankyrin</a> <ul><li><a href="/wiki/Ankyrin-1" title="Ankyrin-1">ANK1</a></li> <li><a href="/wiki/Ankyrin-2" title="Ankyrin-2">ANK2</a></li> <li><a href="/wiki/Ankyrin-3" title="Ankyrin-3">ANK3</a></li></ul></li> <li><a href="/wiki/Spectrin" title="Spectrin">Spectrin</a> <ul><li><a href="/wiki/Spectrin,_alpha_1" title="Spectrin, alpha 1">SPTA1</a></li> <li><a href="/wiki/SPTAN1" title="SPTAN1">SPTAN1</a></li> <li><a href="/wiki/SPTB" title="SPTB">SPTB</a></li> <li><a href="/wiki/SPTBN1" title="SPTBN1">SPTBN1</a></li> <li><a href="/wiki/SPTBN2" title="SPTBN2">SPTBN2</a></li> <li><a href="/wiki/SPTBN4" title="SPTBN4">SPTBN4</a></li> <li><a href="/wiki/SPTBN5" title="SPTBN5">SPTBN5</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Plakin" title="Plakin">Plakins</a> <ul><li><a href="/wiki/Corneodesmosin" title="Corneodesmosin">Corneodesmosin</a></li> <li><a href="/wiki/Desmoplakin" title="Desmoplakin">Desmoplakin</a></li> <li><a href="/wiki/Dystonin" title="Dystonin">Dystonin</a></li> <li><a href="/wiki/Envoplakin" title="Envoplakin">Envoplakin</a></li> <li><a href="/wiki/MACF1" title="MACF1">MACF1</a></li> <li><a href="/wiki/Periplakin" title="Periplakin">Periplakin</a></li> <li><a href="/wiki/Plectin" title="Plectin">Plectin</a></li></ul></li> <li><a href="/wiki/Talin_(protein)" title="Talin (protein)">Talin</a> <ul><li><a href="/wiki/TLN1" title="TLN1">TLN1</a></li></ul></li> <li><a href="/wiki/Vinculin" title="Vinculin">Vinculin</a></li> <li><a href="/wiki/Plakophilin" title="Plakophilin">Plakophilin</a> <ul><li><a href="/wiki/Plakophilin-1" title="Plakophilin-1">PKP1</a></li> <li><a href="/wiki/Plakophilin-2" title="Plakophilin-2">PKP2</a></li></ul></li> <li><a href="/wiki/ACF7" class="mw-redirect" title="ACF7">ACF7</a></li> <li><a href="/wiki/PLEKHA7" title="PLEKHA7">PLEKHA7</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Nonhuman</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/Major_sperm_protein" title="Major sperm protein">Major sperm proteins</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a> <ul><li><a href="/wiki/Crescentin" title="Crescentin">Crescentin</a></li> <li><a href="/wiki/FtsZ" title="FtsZ">FtsZ</a></li> <li><a href="/wiki/MreB" title="MreB">MreB</a></li> <li><a href="/wiki/ParM" title="ParM">ParM</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div><i>See also: <a href="/wiki/Template:Cytoskeletal_defects" title="Template:Cytoskeletal defects">cytoskeletal defects</a></i></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" 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href="https://www.wikidata.org/wiki/Q154626#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></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><span class="uid"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4121944-2">Germany</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Cytoskeleton"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85035256">United States</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Cytosquelette"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb12113172p">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Cytosquelette"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb12113172p">BnF data</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00576209">Japan</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="cytoskelet"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph705648&CON_LNG=ENG">Czech Republic</a></span></span></li><li><span 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