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<span>Types</span> </div> </a> <ul id="toc-Types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mammalian_sphingolipid_metabolism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mammalian_sphingolipid_metabolism"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Mammalian sphingolipid metabolism</span> </div> </a> <ul id="toc-Mammalian_sphingolipid_metabolism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Functions_of_mammalian_sphingolipids" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Functions_of_mammalian_sphingolipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Functions of mammalian sphingolipids</span> </div> </a> <ul id="toc-Functions_of_mammalian_sphingolipids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_sphingolipids" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Other_sphingolipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Other sphingolipids</span> </div> </a> <button aria-controls="toc-Other_sphingolipids-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 Other sphingolipids subsection</span> </button> <ul id="toc-Other_sphingolipids-sublist" class="vector-toc-list"> <li id="toc-Yeast_sphingolipids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Yeast_sphingolipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Yeast sphingolipids</span> </div> </a> <ul id="toc-Yeast_sphingolipids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plant_sphingolipids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plant_sphingolipids"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Plant sphingolipids</span> </div> </a> <ul id="toc-Plant_sphingolipids-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Disorders" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Disorders"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Disorders</span> </div> </a> <ul id="toc-Disorders-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Additional_images" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Additional_images"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Additional images</span> </div> </a> <ul id="toc-Additional_images-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 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">Sphingolipid</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 32 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-32" 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">32 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B4%D8%AD%D9%85%D9%8A%D8%A7%D8%AA_%D8%B3%D9%81%D9%8A%D9%86%D8%AC%D9%88%D9%84%D9%8A%D8%A9" title="شحميات سفينجولية – Arabic" lang="ar" hreflang="ar" data-title="شحميات سفينجولية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%A1%D1%84%D1%96%D0%BD%D0%B3%D0%B0%D0%BB%D1%96%D0%BF%D1%96%D0%B4%D1%8B" title="Сфінгаліпіды – Belarusian" lang="be" hreflang="be" data-title="Сфінгаліпіды" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A1%D1%84%D0%B8%D0%BD%D0%B3%D0%BE%D0%BB%D0%B8%D0%BF%D0%B8%D0%B4" title="Сфинголипид – Bulgarian" lang="bg" hreflang="bg" data-title="Сфинголипид" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Esfingol%C3%ADpid" title="Esfingolípid – Catalan" lang="ca" hreflang="ca" data-title="Esfingolípid" 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/Sfingolipidy" title="Sfingolipidy – Czech" lang="cs" hreflang="cs" data-title="Sfingolipidy" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Sphingolipide" title="Sphingolipide – German" lang="de" hreflang="de" data-title="Sphingolipide" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Esfingol%C3%ADpido" title="Esfingolípido – Spanish" lang="es" hreflang="es" data-title="Esfingolípido" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Esfingolipido" title="Esfingolipido – Basque" lang="eu" hreflang="eu" data-title="Esfingolipido" 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%D9%81%D9%86%DA%AF%D9%88%D9%84%DB%8C%D9%BE%DB%8C%D8%AF" title="اسفنگولیپید – Persian" lang="fa" hreflang="fa" data-title="اسفنگولیپید" 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/Sphingolipide" title="Sphingolipide – French" lang="fr" hreflang="fr" data-title="Sphingolipide" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Esfingol%C3%ADpido" title="Esfingolípido – Galician" lang="gl" hreflang="gl" data-title="Esfingolípido" 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%8A%A4%ED%95%91%EA%B3%A0%EC%A7%80%EC%A7%88" title="스핑고지질 – Korean" lang="ko" hreflang="ko" data-title="스핑고지질" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Sfingolipid" title="Sfingolipid – Indonesian" lang="id" hreflang="id" data-title="Sfingolipid" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Sfingolipidi" title="Sfingolipidi – Italian" lang="it" hreflang="it" data-title="Sfingolipidi" 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%A1%D7%A4%D7%99%D7%A0%D7%92%D7%95%D7%9C%D7%99%D7%A4%D7%99%D7%93" title="ספינגוליפיד – Hebrew" lang="he" hreflang="he" data-title="ספינגוליפיד" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Szfingolipidek" title="Szfingolipidek – Hungarian" lang="hu" hreflang="hu" data-title="Szfingolipidek" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Sfingolipid" title="Sfingolipid – Malay" lang="ms" hreflang="ms" data-title="Sfingolipid" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Sfingolipide" title="Sfingolipide – Dutch" lang="nl" hreflang="nl" data-title="Sfingolipide" 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/%E3%82%B9%E3%83%95%E3%82%A3%E3%83%B3%E3%82%B4%E8%84%82%E8%B3%AA" title="スフィンゴ脂質 – Japanese" lang="ja" hreflang="ja" data-title="スフィンゴ脂質" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Sfingolipidy" title="Sfingolipidy – Polish" lang="pl" hreflang="pl" data-title="Sfingolipidy" 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/Esfingolip%C3%ADdio" title="Esfingolipídio – Portuguese" lang="pt" hreflang="pt" data-title="Esfingolipídio" 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/Sfingolipid%C4%83" title="Sfingolipidă – Romanian" lang="ro" hreflang="ro" data-title="Sfingolipidă" 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%A1%D1%84%D0%B8%D0%BD%D0%B3%D0%BE%D0%BB%D0%B8%D0%BF%D0%B8%D0%B4%D1%8B" title="Сфинголипиды – Russian" lang="ru" hreflang="ru" data-title="Сфинголипиды" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Sfingolipid" title="Sfingolipid – Slovenian" lang="sl" hreflang="sl" data-title="Sfingolipid" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Sfingolipid" title="Sfingolipid – Serbian" lang="sr" hreflang="sr" data-title="Sfingolipid" 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/Sfingolipid" title="Sfingolipid – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Sfingolipid" 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/Sfingolipidi" title="Sfingolipidi – Finnish" lang="fi" hreflang="fi" data-title="Sfingolipidi" 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/Sfingolipid" title="Sfingolipid – Swedish" lang="sv" hreflang="sv" data-title="Sfingolipid" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%AA%E0%B8%9F%E0%B8%B4%E0%B8%87%E0%B9%82%E0%B8%81%E0%B8%A5%E0%B8%B4%E0%B8%9E%E0%B8%B4%E0%B8%94" title="สฟิงโกลิพิด – Thai" lang="th" hreflang="th" data-title="สฟิงโกลิพิด" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D1%84%D1%96%D0%BD%D0%B3%D0%BE%D0%BB%D1%96%D0%BF%D1%96%D0%B4%D0%B8" title="Сфінголіпіди – Ukrainian" lang="uk" hreflang="uk" data-title="Сфінголіпіди" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%8D%85%E4%BA%BA%E8%84%82" title="獅人脂 – Cantonese" lang="yue" hreflang="yue" data-title="獅人脂" 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/%E9%9E%98%E8%84%82" title="鞘脂 – Chinese" lang="zh" hreflang="zh" data-title="鞘脂" 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/Q410395#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div 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class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Family of chemical compounds</div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid 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class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Sphingolipid" title="Special:EditPage/Sphingolipid">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Sphingolipid%22">"Sphingolipid"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Sphingolipid%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Sphingolipid%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Sphingolipid%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Sphingolipid%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Sphingolipid%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">January 2023</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Sphingolipids_general_structures.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Sphingolipids_general_structures.png/500px-Sphingolipids_general_structures.png" decoding="async" width="500" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Sphingolipids_general_structures.png/750px-Sphingolipids_general_structures.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Sphingolipids_general_structures.png/1000px-Sphingolipids_general_structures.png 2x" data-file-width="5186" data-file-height="2277" /></a><figcaption>General structures of sphingolipids</figcaption></figure> <p><b>Sphingolipids</b> are a class of <a href="/wiki/Lipid" title="Lipid">lipids</a> containing a backbone of sphingoid bases, which are a set of <a href="/wiki/Aliphatic" class="mw-redirect" title="Aliphatic">aliphatic</a> <a href="/wiki/Amino" class="mw-redirect" title="Amino">amino</a> <a href="/wiki/Alcohol_(chemistry)" title="Alcohol (chemistry)">alcohols</a> that includes <a href="/wiki/Sphingosine" title="Sphingosine">sphingosine</a>. They were discovered in brain extracts in the 1870s and were named after the mythological <a href="/wiki/Sphinx" title="Sphinx">sphinx</a> because of their enigmatic nature.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These compounds play important roles in <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> and <a href="/wiki/Cell_recognition" class="mw-redirect" title="Cell recognition">cell recognition</a>.<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> <a href="/wiki/Sphingolipidoses" title="Sphingolipidoses">Sphingolipidoses</a>, or disorders of sphingolipid metabolism, have particular impact on <a href="/wiki/Neural_tissue" class="mw-redirect" title="Neural tissue">neural tissue</a>. A sphingolipid with a terminal hydroxyl group is a <a href="/wiki/Ceramide" title="Ceramide">ceramide</a>. Other common groups bonded to the terminal oxygen atom include <a href="/wiki/Phosphocholine" title="Phosphocholine">phosphocholine</a>, yielding a <a href="/wiki/Sphingomyelin" title="Sphingomyelin">sphingomyelin</a>, and various sugar <a href="/wiki/Monomer" title="Monomer">monomers</a> or <a href="/wiki/Dimerization_(chemistry)" class="mw-redirect" title="Dimerization (chemistry)">dimers</a>, yielding <a href="/wiki/Cerebroside" title="Cerebroside">cerebrosides</a> and <a href="/wiki/Globoside" title="Globoside">globosides</a>, respectively. Cerebrosides and globosides are collectively known as <a href="/wiki/Glycosphingolipid" title="Glycosphingolipid">glycosphingolipids</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The long-chain bases, sometimes simply known as sphingoid bases, are the first non-transient products of <i><a href="/wiki/De_novo_synthesis" title="De novo synthesis">de novo</a></i> sphingolipid synthesis in both yeast and mammals. These compounds, specifically known as <a href="/wiki/Phytosphingosine" title="Phytosphingosine">phytosphingosine</a> and <a href="/wiki/Safingol" title="Safingol">dihydrosphingosine</a> (also known as sphinganine,<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> although this term is less common), are mainly C<sub>18</sub> compounds, with somewhat lower levels of C<sub>20</sub> bases.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Ceramides and glycosphingolipids are <i>N</i>-acyl derivatives of these compounds.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The sphingosine backbone is O-linked to a (usually) charged head group such as <a href="/wiki/Ethanolamine" title="Ethanolamine">ethanolamine</a>, <a href="/wiki/Serine" title="Serine">serine</a>, or <a href="/wiki/Choline" title="Choline">choline</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p><p>The backbone is also amide-linked to an <a href="/wiki/Acyl_group" title="Acyl group">acyl group</a>, such as a <a href="/wiki/Fatty_acid" title="Fatty acid">fatty acid</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=2" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Simple sphingolipids, which include the sphingoid bases and ceramides, make up the early products of the sphingolipid synthetic pathways. </p> <ul><li>Sphingoid bases are the fundamental building blocks of all sphingolipids. The main mammalian sphingoid bases are dihydrosphingosine and sphingosine, while dihydrosphingosine and phytosphingosine are the principal sphingoid bases in yeast.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Sphingosine, dihydrosphingosine, and phytosphingosine may be phosphorylated.</li> <li><a href="/wiki/Ceramides" class="mw-redirect" title="Ceramides">Ceramides</a>, as a general class, are <i>N</i>-acylated sphingoid bases lacking additional head groups. <ul><li>Dihydroceramide is produced by <i>N</i>-acylation of dihydrosphingosine. Dihydroceramide is found in both yeast and mammalian systems.</li> <li><a href="/wiki/Ceramide" title="Ceramide">Ceramide</a> is produced in mammalian systems by desaturation of dihydroceramide by dihydroceramide desaturase 1 (DES1). This highly bioactive molecule may also be phosphorylated to form <a href="/w/index.php?title=Ceramide-1-phosphate&action=edit&redlink=1" class="new" title="Ceramide-1-phosphate (page does not exist)">ceramide-1-phosphate</a>.</li> <li><a href="/wiki/Phytoceramide" class="mw-redirect" title="Phytoceramide">Phytoceramide</a> is produced in yeast by hydroxylation of dihydroceramide at C-4.</li></ul></li></ul> <p>Complex sphingolipids may be formed by addition of head groups to ceramide or phytoceramide: </p> <ul><li><a href="/wiki/Sphingomyelin" title="Sphingomyelin">Sphingomyelins</a> have a <a href="/wiki/Phosphocholine" title="Phosphocholine">phosphocholine</a> or <a href="/wiki/Phosphoethanolamine" class="mw-redirect" title="Phosphoethanolamine">phosphoethanolamine</a> molecule with an <a href="/wiki/Ester" title="Ester">ester linkage</a> to the 1-hydroxy group of a ceramide.</li> <li><a href="/wiki/Glycosphingolipid" title="Glycosphingolipid">Glycosphingolipids</a> are ceramides with one or more <a href="/wiki/Sugar" title="Sugar">sugar</a> residues joined in a β-<a href="/wiki/Glycosidic" class="mw-redirect" title="Glycosidic">glycosidic</a> linkage at the 1-hydroxyl position (see image). <ul><li><a href="/wiki/Cerebroside" title="Cerebroside">Cerebrosides</a> have a single <a href="/wiki/Glucose" title="Glucose">glucose</a> or <a href="/wiki/Galactose" title="Galactose">galactose</a> at the 1-hydroxy position. <ul><li><a href="/wiki/Sulfatide" title="Sulfatide">Sulfatides</a> are sulfated cerebrosides.</li></ul></li> <li><a href="/wiki/Gangliosides" class="mw-redirect" title="Gangliosides">Gangliosides</a> have at least three sugars, one of which must be <a href="/wiki/Sialic_acid" title="Sialic acid">sialic acid</a>.</li></ul></li> <li><a href="/wiki/Inositol" title="Inositol">Inositol</a>-containing ceramides, which are derived from phytoceramide, are produced in yeast. These include inositol phosphorylceramide, <a href="/wiki/Mannose" title="Mannose">mannose</a> inositol phosphorylceramide, and mannose diinositol phosphorylceramide.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Mammalian_sphingolipid_metabolism">Mammalian sphingolipid metabolism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=3" title="Edit section: Mammalian sphingolipid metabolism"><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">See also: <a href="/wiki/Palmitoyl-CoA#Sphingolipid_biosynthesis" title="Palmitoyl-CoA">Palmitoyl-CoA § Sphingolipid biosynthesis</a></div> <p><i>De novo</i> sphingolipid synthesis begins with formation of 3-keto-dihydrosphingosine by <a href="/wiki/Serine_palmitoyltransferase" class="mw-redirect" title="Serine palmitoyltransferase">serine palmitoyltransferase</a>.<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> The preferred substrates for this reaction are <a href="/wiki/Palmitoyl-CoA" title="Palmitoyl-CoA">palmitoyl-CoA</a> and <a href="/wiki/Serine" title="Serine">serine</a>. However, studies have demonstrated that serine palmitoyltransferase has some activity toward other species of <a href="/wiki/Fatty_acyl-CoA" class="mw-redirect" title="Fatty acyl-CoA">fatty acyl-CoA</a><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> and alternative <a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">amino acids</a>,<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> and the diversity of sphingoid bases has recently been reviewed.<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> Next, 3-keto-dihydrosphingosine is reduced to form dihydrosphingosine. Dihydrosphingosine is acylated by one of six (dihydro)-ceramide synthase, <a href="/w/index.php?title=CerS&action=edit&redlink=1" class="new" title="CerS (page does not exist)">CerS</a> - originally termed LASS - to form dihydroceramide.<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> The six CerS enzymes have different specificity for <a href="/wiki/Acyl-CoA" title="Acyl-CoA">acyl-CoA</a> substrates, resulting in the generation of dihydroceramides with differing chain lengths (ranging from C14-C26). Dihydroceramides are then desaturated to form ceramide.<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> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Sphingolipidoses.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2a/Sphingolipidoses.svg/400px-Sphingolipidoses.svg.png" decoding="async" width="400" height="335" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2a/Sphingolipidoses.svg/600px-Sphingolipidoses.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2a/Sphingolipidoses.svg/800px-Sphingolipidoses.svg.png 2x" data-file-width="1238" data-file-height="1036" /></a><figcaption>Metabolic pathways of various forms of sphingolipids. <a href="/wiki/Sphingolipidoses" title="Sphingolipidoses">Sphingolipidoses</a> are labeled at corresponding stages that are deficient.</figcaption></figure> <p>De novo generated <a href="/wiki/Ceramide" title="Ceramide">ceramide</a> is the central hub of the sphingolipid network and subsequently has several fates. It may be phosphorylated by <a href="/wiki/Ceramide_kinase" title="Ceramide kinase">ceramide kinase</a> to form ceramide-1-phosphate. Alternatively, it may be glycosylated by <a href="/wiki/Glucosylceramide_synthase" class="mw-redirect" title="Glucosylceramide synthase">glucosylceramide synthase</a> or <a href="/w/index.php?title=Galactosylceramide_synthase&action=edit&redlink=1" class="new" title="Galactosylceramide synthase (page does not exist)">galactosylceramide synthase</a>. Additionally, it can be converted to <a href="/wiki/Sphingomyelin" title="Sphingomyelin">sphingomyelin</a> by the addition of a <a href="/wiki/Phosphorylcholine" title="Phosphorylcholine">phosphorylcholine</a> headgroup by <a href="/wiki/Sphingomyelin_synthase" title="Sphingomyelin synthase">sphingomyelin synthase</a>. <a href="/wiki/Diacylglycerol" class="mw-redirect" title="Diacylglycerol">Diacylglycerol</a> is generated by this process. Finally, ceramide may be broken down by a <a href="/wiki/Ceramidase" title="Ceramidase">ceramidase</a> to form <a href="/wiki/Sphingosine" title="Sphingosine">sphingosine</a>. <a href="/wiki/Sphingosine" title="Sphingosine">Sphingosine</a> may be phosphorylated to form sphingosine-1-phosphate. This may be dephosphorylated to reform sphingosine.<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><p>Breakdown pathways allow the reversion of these metabolites to ceramide. The complex glycosphingolipids are hydrolyzed to glucosylceramide and galactosylceramide. These lipids are then hydrolyzed by beta-glucosidases and beta-galactosidases to regenerate ceramide. Similarly, sphingomyelin may be broken down by sphingomyelinase to form ceramide.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p><p>The only route by which sphingolipids are converted to non-sphingolipids is through sphingosine-1-phosphate lyase. This forms ethanolamine phosphate and hexadecenal.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Functions_of_mammalian_sphingolipids">Functions of mammalian sphingolipids</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=4" title="Edit section: Functions of mammalian sphingolipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sphingolipids are commonly believed to protect the cell surface against harmful environmental factors by forming a mechanically stable and chemically resistant outer leaflet of the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> <a href="/wiki/Lipid_bilayer" title="Lipid bilayer">lipid bilayer</a>. Certain complex <a href="/wiki/Glycosphingolipids" class="mw-redirect" title="Glycosphingolipids">glycosphingolipids</a> were found to be involved in specific functions, such as <a href="/wiki/Cell_signaling" title="Cell signaling">cell recognition and signaling</a>. Cell recognition depends mainly on the physical properties of the sphingolipids, whereas signaling involves specific interactions of the glycan structures of glycosphingolipids with similar lipids present on neighboring cells or with <a href="/wiki/Protein" title="Protein">proteins</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p><p>Recently, simple sphingolipid <a href="/wiki/Metabolite" title="Metabolite">metabolites</a>, such as ceramide and <a href="/wiki/Sphingosine-1-phosphate" title="Sphingosine-1-phosphate">sphingosine-1-phosphate</a>, have been shown to be important mediators in the signaling cascades involved in <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, <a href="/wiki/Cell_growth" title="Cell growth">proliferation</a>, stress responses, <a href="/wiki/Necrosis" title="Necrosis">necrosis</a>, <a href="/wiki/Inflammation" title="Inflammation">inflammation</a>, <a href="/wiki/Autophagy" title="Autophagy">autophagy</a>, <a href="/wiki/Senescence" title="Senescence">senescence</a>, and <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><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> Ceramide-based lipids self-aggregate in <a href="/wiki/Cell_membrane" title="Cell membrane">cell membranes</a> and form separate <a href="/wiki/Phase_(matter)" title="Phase (matter)">phases</a> less fluid than the bulk phospholipids. These sphingolipid-based microdomains, or "<a href="/wiki/Lipid_raft" title="Lipid raft">lipid rafts</a>" were originally proposed to sort membrane proteins along the cellular pathways of membrane transport. At present, most research focuses on the organizing function during signal transduction.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>Sphingolipids are synthesized in a pathway that begins in the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">ER</a> and is completed in the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a>, but these lipids are enriched in the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> and in <a href="/wiki/Endosomes" class="mw-redirect" title="Endosomes">endosomes</a>, where they perform many of their functions.<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> Transport occurs via vesicles and monomeric transport in the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>. Sphingolipids are virtually absent from <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> and the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">ER</a>, but constitute a 20-35 molar fraction of plasma membrane lipids.<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> </p><p>In experimental animals, feeding sphingolipids inhibits <a href="/wiki/Colorectal_cancer" title="Colorectal cancer">colon carcinogenesis</a>, reduces <a href="/wiki/Low-density_lipoprotein" title="Low-density lipoprotein">LDL cholesterol</a> and elevates <a href="/wiki/High-density_lipoprotein" title="High-density lipoprotein">HDL cholesterol</a>.<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> <div class="mw-heading mw-heading2"><h2 id="Other_sphingolipids">Other sphingolipids</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=5" title="Edit section: Other sphingolipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sphingolipids are universal in <a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a> but are rare in <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>, meaning that they are evolutionally very old. Bacteria that do produce sphingolipids are found in some members of the <a href="/wiki/Superphylum" class="mw-redirect" title="Superphylum">superphylum</a> <a href="/wiki/FCB_group" class="mw-redirect" title="FCB group">FCB group</a> (<a href="/wiki/Sphingobacteria_(phylum)" class="mw-redirect" title="Sphingobacteria (phylum)">Sphingobacteria</a>), particularly family <a href="/wiki/Sphingomonadaceae" title="Sphingomonadaceae">Sphingomonadaceae</a>, some members of the <a href="/wiki/Bdellovibrionota" title="Bdellovibrionota">Bdellovibrionota</a>, and some members of the <a href="/wiki/Myxococcota" title="Myxococcota">Myxococcota</a>.<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> <div class="mw-heading mw-heading3"><h3 id="Yeast_sphingolipids">Yeast sphingolipids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=6" title="Edit section: Yeast sphingolipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Because of the incredible complexity of mammalian systems, yeast are often used as a <a href="/wiki/Model_organism" title="Model organism">model organism</a> for working out new pathways. These single-celled organisms are often more genetically tractable than mammalian cells, and strain libraries are available to supply strains harboring almost any non-lethal <a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frame</a> single deletion. The two most commonly used yeasts are <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> and <i><a href="/wiki/Schizosaccharomyces_pombe" title="Schizosaccharomyces pombe">Schizosaccharomyces pombe</a></i>, although research is also done in the pathogenic yeast <i><a href="/wiki/Candida_albicans" title="Candida albicans">Candida albicans</a></i>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p><p>In addition to the important structural functions of complex sphingolipids (inositol phosphorylceramide and its mannosylated derivatives), the sphingoid bases <a href="/wiki/Phytosphingosine" title="Phytosphingosine">phytosphingosine</a> and dihydrosphingosine (sphinganine) play vital signaling roles in <i>S. cerevisiae</i>. These effects include regulation of <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>, ubiquitin-dependent <a href="/wiki/Proteolysis" title="Proteolysis">proteolysis</a> (and, thus, regulation of nutrient uptake <sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup>), <a href="/wiki/Cytoskeletal" class="mw-redirect" title="Cytoskeletal">cytoskeletal</a> dynamics, the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>, <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, posttranslational protein modification, and the heat stress response.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Additionally, modulation of sphingolipid metabolism by <a href="/wiki/Phosphatidylinositol_(4,5)-bisphosphate" class="mw-redirect" title="Phosphatidylinositol (4,5)-bisphosphate">phosphatidylinositol (4,5)-bisphosphate</a> signaling <i>via</i> Slm1p and Slm2p and <a href="/wiki/Calcineurin" title="Calcineurin">calcineurin</a> has recently been described.<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> Additionally, a substrate-level interaction has been shown between complex sphingolipid synthesis and cycling of <a href="/wiki/Phosphatidylinositol_4-phosphate" title="Phosphatidylinositol 4-phosphate">phosphatidylinositol 4-phosphate</a> by the phosphatidylinositol kinase Stt4p and the lipid phosphatase Sac1p.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Plant_sphingolipids">Plant sphingolipids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=7" title="Edit section: Plant sphingolipids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Higher plants contain a wider variety of sphingolipids than animals and fungi.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Disorders">Disorders</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=8" title="Edit section: Disorders"><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/Sphingolipidoses" title="Sphingolipidoses">Sphingolipidoses</a></div> <p>There are several disorders of sphingolipid metabolism, known as <a href="/wiki/Sphingolipidoses" title="Sphingolipidoses">sphingolipidoses</a>. The main members of this group are <a href="/wiki/Niemann-Pick_disease" class="mw-redirect" title="Niemann-Pick disease">Niemann-Pick disease</a>, <a href="/wiki/Fabry_disease" title="Fabry disease">Fabry disease</a>, <a href="/wiki/Krabbe_disease" title="Krabbe disease">Krabbe disease</a>, <a href="/wiki/Gaucher_disease" class="mw-redirect" title="Gaucher disease">Gaucher disease</a>, <a href="/wiki/Tay%E2%80%93Sachs_disease" title="Tay–Sachs disease">Tay–Sachs disease</a> and <a href="/wiki/Metachromatic_leukodystrophy" title="Metachromatic leukodystrophy">Metachromatic leukodystrophy</a>. They are generally inherited in an <a href="/wiki/Autosomal_recessive" class="mw-redirect" title="Autosomal recessive">autosomal recessive</a> fashion, but notably <a href="/wiki/Fabry_disease" title="Fabry disease">Fabry disease</a> is <a href="/wiki/X-linked" class="mw-redirect" title="X-linked">X-linked</a>. Taken together, sphingolipidoses have an <a href="/wiki/Incidence_(epidemiology)" title="Incidence (epidemiology)">incidence</a> of approximately 1 in 10,000, but substantially more in certain populations such as <a href="/wiki/Ashkenazi_Jews" title="Ashkenazi Jews">Ashkenazi Jews</a>. <a href="/wiki/Enzyme_replacement_therapy" title="Enzyme replacement therapy">Enzyme replacement therapy</a> is available to treat mainly <a href="/wiki/Fabry_disease" title="Fabry disease">Fabry disease</a> and <a href="/wiki/Gaucher_disease" class="mw-redirect" title="Gaucher disease">Gaucher disease</a>, and people with these types of sphingolipidoses may live well into adulthood. The other types are generally fatal by age 1 to 5 years for infantile forms, but progression may be mild for juvenile- or adult-onset forms.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2017)">citation needed</span></a></i>]</sup> </p><p>Sphingolipids have also been implicated with the frataxin protein (Fxn), the deficiency of which is associated with <a href="/wiki/Friedreich%27s_ataxia" title="Friedreich's ataxia">Friedreich's ataxia</a> (FRDA). Loss of Fxn in the nervous system in mice also activates an iron/sphingolipid/PDK1/Mef2 pathway, indicating that the mechanism is evolutionarily conserved. Furthermore, sphingolipid levels and PDK1 activity are also increased in hearts of FRDA patients, suggesting that a similar pathway is affected in FRDA.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Other research has demonstrated that iron accumulation in the nervous systems of flies enhances the synthesis of sphingolipids, which in turn activates 3-phosphoinositide dependent protein kinase-1 (Pdk1) and myocyte enhancer factor-2 (Mef2) to trigger neurodegeneration of adult photoreceptors.<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><p>Sphingolipids play a key role in neuronal survival in Parkinson's Disease (PD) and their catabolic pathway alteration in the brain is partly represented in cerebrospinal fluid and blood tissues (Table1) and have the diagnostic potential.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Additional_images">Additional images</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=9" title="Edit section: Additional images"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Sphingosine_structure.svg" class="mw-file-description" title="Sphingosine"><img alt="Sphingosine" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Sphingosine_structure.svg/120px-Sphingosine_structure.svg.png" decoding="async" width="120" height="24" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Sphingosine_structure.svg/180px-Sphingosine_structure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Sphingosine_structure.svg/240px-Sphingosine_structure.svg.png 2x" data-file-width="512" data-file-height="101" /></a></span></div> <div class="gallerytext"><a href="/wiki/Sphingosine" title="Sphingosine">Sphingosine</a></div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=10" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Sphingosyl_phosphatide" title="Sphingosyl phosphatide">Sphingosyl phosphatide</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Sphingolipid&action=edit&section=11" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFChunHartung2010" class="citation journal cs1">Chun J, Hartung HP (2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2859693">"Mechanism of action of oral fingolimod (FTY720) in multiple sclerosis"</a>. <i>Clinical Neuropharmacology</i>. <b>33</b> (2): 91–101. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1097%2Fwnf.0b013e3181cbf825">10.1097/wnf.0b013e3181cbf825</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/PMC2859693">2859693</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/20061941">20061941</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Clinical+Neuropharmacology&rft.atitle=Mechanism+of+action+of+oral+fingolimod+%28FTY720%29+in+multiple+sclerosis&rft.volume=33&rft.issue=2&rft.pages=91-101&rft.date=2010&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2859693%23id-name%3DPMC&rft_id=info%3Apmid%2F20061941&rft_id=info%3Adoi%2F10.1097%2Fwnf.0b013e3181cbf825&rft.aulast=Chun&rft.aufirst=J&rft.au=Hartung%2C+HP&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2859693&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchnaarSandhoffTiemeyerKinoshita2022" class="citation cs2">Schnaar, Ronald L.; Sandhoff, Roger; Tiemeyer, Michael; 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Molecular Cell Biology</i>. <b>19</b> (5): 281–296. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrm.2017.138">10.1038/nrm.2017.138</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1471-0080">1471-0080</a>. <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/29410529">29410529</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews.+Molecular+Cell+Biology&rft.atitle=Understanding+the+diversity+of+membrane+lipid+composition&rft.volume=19&rft.issue=5&rft.pages=281-296&rft.date=2018-05&rft.issn=1471-0080&rft_id=info%3Apmid%2F29410529&rft_id=info%3Adoi%2F10.1038%2Fnrm.2017.138&rft.aulast=Harayama&rft.aufirst=Takeshi&rft.au=Riezman%2C+Howard&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F29410529%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://www.sigmaaldrich.com/US/en/product/sigma/d3314">|SIGMA&N5=SEARCH_CONCAT_PNO|BRAND_KEY&F=SPEC Product page at Sigma Aldrich</a></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDickson1998" class="citation journal cs1">Dickson RC (1998). <a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.biochem.67.1.27">"Sphingolipid functions in Saccharomyces cerevisiae: comparison to mammals"</a>. <i>Annual Review of Biochemistry</i>. <b>67</b>: 27–48. <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/10.1146%2Fannurev.biochem.67.1.27">10.1146/annurev.biochem.67.1.27</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/9759481">9759481</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Biochemistry&rft.atitle=Sphingolipid+functions+in+Saccharomyces+cerevisiae%3A+comparison+to+mammals&rft.volume=67&rft.pages=27-48&rft.date=1998&rft_id=info%3Adoi%2F10.1146%2Fannurev.biochem.67.1.27&rft_id=info%3Apmid%2F9759481&rft.aulast=Dickson&rft.aufirst=RC&rft_id=https%3A%2F%2Fdoi.org%2F10.1146%252Fannurev.biochem.67.1.27&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text">A brief, very comprehensible review is given in Gunstone, F. 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(identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27901468">27901468</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=eLife&rft.atitle=Loss+of+Frataxin+activates+the+iron%2Fsphingolipid%2FPDK1%2FMef2+pathway+in+mammals&rft.volume=5&rft.date=2016-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5130293%23id-name%3DPMC&rft_id=info%3Apmid%2F27901468&rft_id=info%3Adoi%2F10.7554%2FeLife.20732&rft.aulast=Chen&rft.aufirst=K&rft.au=Ho%2C+TS&rft.au=Lin%2C+G&rft.au=Tan%2C+KL&rft.au=Rasband%2C+MN&rft.au=Bellen%2C+HJ&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5130293&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" class="Z3988"></span></span> </li> <li id="cite_note-32"><span class="mw-cite-backlink"><b><a href="#cite_ref-32">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChenLinHaeltermanHo2016" class="citation journal cs1">Chen K, Lin G, Haelterman NA, Ho TS, Li T, Li Z, Duraine L, Graham BH, Jaiswal M, Yamamoto S, Rasband MN, Bellen HJ (June 2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4956409">"Loss of Frataxin induces iron toxicity, sphingolipid synthesis, and Pdk1/Mef2 activation, leading to neurodegeneration"</a>. <i>eLife</i>. <b>5</b>. <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/10.7554%2FeLife.16043">10.7554/eLife.16043</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/PMC4956409">4956409</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/27343351">27343351</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=eLife&rft.atitle=Loss+of+Frataxin+induces+iron+toxicity%2C+sphingolipid+synthesis%2C+and+Pdk1%2FMef2+activation%2C+leading+to+neurodegeneration&rft.volume=5&rft.date=2016-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4956409%23id-name%3DPMC&rft_id=info%3Apmid%2F27343351&rft_id=info%3Adoi%2F10.7554%2FeLife.16043&rft.aulast=Chen&rft.aufirst=K&rft.au=Lin%2C+G&rft.au=Haelterman%2C+NA&rft.au=Ho%2C+TS&rft.au=Li%2C+T&rft.au=Li%2C+Z&rft.au=Duraine%2C+L&rft.au=Graham%2C+BH&rft.au=Jaiswal%2C+M&rft.au=Yamamoto%2C+S&rft.au=Rasband%2C+MN&rft.au=Bellen%2C+HJ&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4956409&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" class="Z3988"></span></span> </li> <li id="cite_note-33"><span class="mw-cite-backlink"><b><a href="#cite_ref-33">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEsfandiaryFinkelsteinVoelckerRudd2022" class="citation journal cs1">Esfandiary, Ali; Finkelstein, David Isaac; Voelcker, Nicolas Hans; Rudd, David (2022-04-15). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9028315">"Clinical Sphingolipids Pathway in Parkinson's Disease: From GCase to Integrated-Biomarker Discovery"</a>. <i>Cells</i>. <b>11</b> (8): 1353. <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/10.3390%2Fcells11081353">10.3390/cells11081353</a></span>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2073-4409">2073-4409</a>. <a href="/wiki/PMC_(identifier)" 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cells&rft.atitle=Clinical+Sphingolipids+Pathway+in+Parkinson%27s+Disease%3A+From+GCase+to+Integrated-Biomarker+Discovery&rft.volume=11&rft.issue=8&rft.pages=1353&rft.date=2022-04-15&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC9028315%23id-name%3DPMC&rft.issn=2073-4409&rft_id=info%3Apmid%2F35456032&rft_id=info%3Adoi%2F10.3390%2Fcells11081353&rft.aulast=Esfandiary&rft.aufirst=Ali&rft.au=Finkelstein%2C+David+Isaac&rft.au=Voelcker%2C+Nicolas+Hans&rft.au=Rudd%2C+David&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC9028315&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASphingolipid" 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=Sphingolipid&action=edit&section=12" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Sphingolipids">Sphingolipids</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLaurila2022" class="citation journal cs1">Laurila, Pirkka-Pekka; et al. (December 2022). <span class="id-lock-subscription" title="Paid subscription required"><a rel="nofollow" class="external text" href="https://www.nature.com/articles/s43587-022-00309-6">"Sphingolipids accumulate in aged muscle, and their reduction counteracts sarcopenia"</a></span>. <i>Nature Aging</i>. <b>2</b> (12): 1159–1175. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs43587-022-00309-6">10.1038/s43587-022-00309-6</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2662-8465">2662-8465</a>. <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/37118545">37118545</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID 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title="Myelin">Myelin sheath</a></li> <li><a href="/wiki/Node_of_Ranvier" title="Node of Ranvier">Nodes of Ranvier</a></li> <li><a href="/wiki/Nuclear_envelope" title="Nuclear envelope">Nuclear envelope</a></li> <li><a href="/wiki/Phycobilisome" title="Phycobilisome">Phycobilisomes</a></li> <li><a href="/wiki/Porosome" title="Porosome">Porosomes</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="Glycoconjugates,_lipids,_and_glycolipids:_sphingolipids_and_glycosphingolipids,_and_metabolic_intermediates" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link 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title="Glycolipid">glycolipids</a>: <a class="mw-selflink selflink">sphingolipids</a> and <a href="/wiki/Glycosphingolipid" title="Glycosphingolipid">glycosphingolipids</a>, and <a href="/wiki/Metabolic_intermediate" title="Metabolic intermediate">metabolic intermediates</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Simple glycosphingolipids</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cerebroside" title="Cerebroside">Cerebroside</a> <ul><li><a href="/wiki/Galactocerebroside" class="mw-redirect" title="Galactocerebroside">Galactocerebroside</a></li> <li><a href="/wiki/Glucocerebroside" title="Glucocerebroside">Glucocerebroside</a></li></ul></li> <li><a href="/wiki/Sulfatide" title="Sulfatide">Sulfatide</a></li> <li><a href="/wiki/Lactosylceramide" title="Lactosylceramide">Lactosylceramide</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Globosides" class="mw-redirect" title="Globosides">Globosides</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Globotriaosylceramide" title="Globotriaosylceramide">Globotriaosylceramide</a></li> <li><a href="/wiki/Stage-specific_embryonic_antigen_3" title="Stage-specific embryonic antigen 3">Stage-specific embryonic antigen 3</a></li> <li><a href="/wiki/Globo_H" title="Globo H">Globo H</a></li> <li><a href="/wiki/Forssman_antigen" title="Forssman antigen">Forssman antigen</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Gangliosides" class="mw-redirect" title="Gangliosides">Gangliosides</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/GM1" title="GM1">GM1</a></li> <li><a href="/wiki/GM2_(ganglioside)" title="GM2 (ganglioside)">GM2</a></li> <li><a href="/wiki/GM3" title="GM3">GM3</a></li> <li><a href="/wiki/GD2" title="GD2">GD2</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 hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sphingomyelin" title="Sphingomyelin">Sphingomyelin</a></li> <li><a href="/wiki/Sphingosine-1-phosphate" title="Sphingosine-1-phosphate">Sphingosine-1-phosphate</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="Metabolism,_lipid_metabolism,_glycolipid_enzymes" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><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:Glycolipid_and_sphingolipid_metabolism_enzymes" title="Template:Glycolipid and sphingolipid metabolism enzymes"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Glycolipid_and_sphingolipid_metabolism_enzymes" title="Template talk:Glycolipid and sphingolipid metabolism enzymes"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Glycolipid_and_sphingolipid_metabolism_enzymes" title="Special:EditPage/Template:Glycolipid and sphingolipid metabolism enzymes"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Metabolism,_lipid_metabolism,_glycolipid_enzymes" style="font-size:114%;margin:0 4em"><a href="/wiki/Metabolism" title="Metabolism">Metabolism</a>, <a href="/wiki/Lipid_metabolism" title="Lipid metabolism">lipid metabolism</a>, <a href="/wiki/Glycolipid" title="Glycolipid">glycolipid</a> <a href="/wiki/Enzyme" title="Enzyme">enzymes</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Sphingolipid</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%">To <a href="/wiki/Glycosphingolipid" title="Glycosphingolipid">glycosphingolipid</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/Glycosyltransferase" title="Glycosyltransferase">Glycosyltransferase</a></li> <li><a href="/wiki/Sulfotransferase" title="Sulfotransferase">Sulfotransferase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">To <a href="/wiki/Ceramide" title="Ceramide">ceramide</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><span class="nobold">From <a href="/wiki/Ganglioside" title="Ganglioside">ganglioside</a></span></dt> <dd><a href="/wiki/Beta-galactosidase" class="mw-redirect" title="Beta-galactosidase">Beta-galactosidase</a></dd> <dd><a href="/wiki/HEXA" title="HEXA">Hexosaminidase A</a></dd> <dd><a href="/wiki/Neuraminidase" title="Neuraminidase">Neuraminidase</a></dd> <dd><a href="/wiki/Glucocerebrosidase" title="Glucocerebrosidase">Glucocerebrosidase</a></dd></dl> <dl><dt><span class="nobold">From <a href="/wiki/Globoside" title="Globoside">globoside</a></span></dt> <dd><a href="/wiki/HEXB" title="HEXB">Hexosaminidase B</a></dd> <dd><a href="/wiki/Alpha-galactosidase" class="mw-redirect" title="Alpha-galactosidase">Alpha-galactosidase</a></dd> <dd><a href="/wiki/Beta-galactosidase" class="mw-redirect" title="Beta-galactosidase">Beta-galactosidase</a></dd> <dd><a href="/wiki/Glucocerebrosidase" title="Glucocerebrosidase">Glucocerebrosidase</a></dd></dl> <dl><dt><span class="nobold">From <a href="/wiki/Sphingomyelin" title="Sphingomyelin">sphingomyelin</a></span></dt> <dd><a href="/wiki/Sphingomyelin_phosphodiesterase" title="Sphingomyelin phosphodiesterase">Sphingomyelin phosphodiesterase</a> <dl><dd><a href="/wiki/Sphingomyelin_phosphodiesterase_1" title="Sphingomyelin phosphodiesterase 1">Sphingomyelin phosphodiesterase 1</a></dd></dl></dd></dl> <dl><dt><span class="nobold">From <a href="/wiki/Sulfatide" title="Sulfatide">sulfatide</a></span></dt> <dd><a href="/wiki/Arylsulfatase_A" title="Arylsulfatase A">Arylsulfatase A</a></dd> <dd><a href="/wiki/Galactosylceramidase" title="Galactosylceramidase">Galactosylceramidase</a></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">To <a href="/wiki/Sphingosine" title="Sphingosine">sphingosine</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/Ceramidase" title="Ceramidase">Ceramidase</a> <ul><li><a href="/wiki/ACER1" title="ACER1">ACER1</a></li> <li><a href="/wiki/ACER2" title="ACER2">ACER2</a></li> <li><a href="/wiki/ACER3" title="ACER3">ACER3</a></li> <li><a href="/wiki/ASAH1" title="ASAH1">ASAH1</a></li> <li><a href="/wiki/ASAH2" title="ASAH2">ASAH2</a></li> <li><a href="/wiki/ASAH2B" title="ASAH2B">ASAH2B</a></li> <li><a href="/wiki/ASAH2C" title="ASAH2C">ASAH2C</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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sphingosine_kinase" title="Sphingosine kinase">Sphingosine kinase</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/Neuronal_ceroid_lipofuscinosis" title="Neuronal ceroid lipofuscinosis">NCL</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/Palmitoyl_protein_thioesterase" class="mw-redirect" title="Palmitoyl protein thioesterase">Palmitoyl protein thioesterase</a></li> <li><a href="/wiki/Tripeptidyl_peptidase_I" title="Tripeptidyl peptidase I">Tripeptidyl peptidase I</a></li> <li><a href="/wiki/CLN3" class="mw-redirect" title="CLN3">CLN3</a></li> <li><a href="/wiki/CLN5" title="CLN5">CLN5</a></li> <li><a href="/wiki/CLN6" title="CLN6">CLN6</a></li> <li><a href="/wiki/CLN8" title="CLN8">CLN8</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ceramide#Pathways_for_ceramide_synthesis" title="Ceramide">Ceramide synthesis</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/Serine_C-palmitoyltransferase" title="Serine C-palmitoyltransferase">Serine C-palmitoyltransferase</a> (<a href="/wiki/SPTLC1" title="SPTLC1">SPTLC1</a>)</li> <li><a href="/wiki/Ceramide_glucosyltransferase" title="Ceramide glucosyltransferase">Ceramide glucosyltransferase</a> (<a href="/wiki/UGCG" title="UGCG">UGCG</a>)</li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output 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