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id="toc-Plasma_membrane_to/from_early_endosomes_(via_recycling_endosomes)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plasma_membrane_to/from_early_endosomes_(via_recycling_endosomes)"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Plasma membrane to/from early endosomes (via recycling endosomes)</span> </div> </a> <ul id="toc-Plasma_membrane_to/from_early_endosomes_(via_recycling_endosomes)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Late_endosomes_to_lysosomes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Late_endosomes_to_lysosomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Late endosomes to lysosomes</span> </div> </a> <ul id="toc-Late_endosomes_to_lysosomes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</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">5</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">6</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" title="Table of Contents" > <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">Endosome</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 29 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-29" 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">29 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%AC%D8%B3%D9%8A%D9%85_%D8%AF%D8%A7%D8%AE%D9%84%D9%8A" title="جسيم داخلي – Arabic" lang="ar" hreflang="ar" data-title="جسيم داخلي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Endosom" title="Endosom – Bosnian" lang="bs" hreflang="bs" data-title="Endosom" 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/Endosoma" title="Endosoma – Catalan" lang="ca" hreflang="ca" data-title="Endosoma" 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/Endozom" title="Endozom – Czech" lang="cs" hreflang="cs" data-title="Endozom" 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/Endosom" title="Endosom – German" lang="de" hreflang="de" data-title="Endosom" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%95%CE%BD%CE%B4%CF%8C%CF%83%CF%89%CE%BC%CE%B1" title="Ενδόσωμα – Greek" lang="el" hreflang="el" data-title="Ενδόσωμα" 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/Endosoma" title="Endosoma – Spanish" lang="es" hreflang="es" data-title="Endosoma" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A7%D9%86%D8%AF%D9%88%D8%B2%D9%88%D9%85" title="اندوزوم – Persian" lang="fa" hreflang="fa" data-title="اندوزوم" 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/Endosome" title="Endosome – French" lang="fr" hreflang="fr" data-title="Endosome" 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/Endosoma" title="Endosoma – Galician" lang="gl" hreflang="gl" data-title="Endosoma" 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%97%94%EB%8F%84%EC%86%9C" title="엔도솜 – Korean" lang="ko" hreflang="ko" data-title="엔도솜" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Endosoma" title="Endosoma – Italian" lang="it" hreflang="it" data-title="Endosoma" 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%90%D7%A0%D7%93%D7%95%D7%96%D7%95%D7%9D" title="אנדוזום – Hebrew" lang="he" hreflang="he" data-title="אנדוזום" 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/Endosz%C3%B3ma" title="Endoszóma – Hungarian" lang="hu" hreflang="hu" data-title="Endoszóma" 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%95%D0%BD%D0%B4%D0%BE%D0%B7%D0%BE%D0%BC" title="Ендозом – Macedonian" lang="mk" hreflang="mk" data-title="Ендозом" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%A8%E3%83%B3%E3%83%89%E3%82%BD%E3%83%BC%E3%83%A0" title="エンドソーム – Japanese" lang="ja" hreflang="ja" data-title="エンドソーム" 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-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Endosomalar" title="Endosomalar – Uzbek" lang="uz" hreflang="uz" data-title="Endosomalar" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Endosomy" title="Endosomy – Polish" lang="pl" hreflang="pl" data-title="Endosomy" 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/Endossomo" title="Endossomo – Portuguese" lang="pt" hreflang="pt" data-title="Endossomo" 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/Endozom" title="Endozom – Romanian" lang="ro" hreflang="ro" data-title="Endozom" 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%AD%D0%BD%D0%B4%D0%BE%D1%81%D0%BE%D0%BC%D0%B0" title="Эндосома – Russian" lang="ru" hreflang="ru" data-title="Эндосома" 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/Endosom" title="Endosom – Slovenian" lang="sl" hreflang="sl" data-title="Endosom" 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/Endozom" title="Endozom – Serbian" lang="sr" hreflang="sr" data-title="Endozom" 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/Endozom" title="Endozom – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Endozom" 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-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Endosom" title="Endosom – Swedish" lang="sv" hreflang="sv" data-title="Endosom" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%80%E0%B8%AD%E0%B8%99%E0%B9%82%E0%B8%94%E0%B9%82%E0%B8%8B%E0%B8%A1" title="เอนโดโซม – Thai" lang="th" hreflang="th" data-title="เอนโดโซม" 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/Endozom" title="Endozom – Turkish" lang="tr" hreflang="tr" data-title="Endozom" 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%95%D0%BD%D0%B4%D0%BE%D1%81%D0%BE%D0%BC%D0%B0" title="Ендосома – Ukrainian" lang="uk" hreflang="uk" data-title="Ендосома" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%83%9E%E5%85%A7%E9%AB%94" title="胞內體 – Chinese" lang="zh" hreflang="zh" data-title="胞內體" data-language-autonym="中文" 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free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Vacuole to which materials ingested by endocytosis are delivered</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">"Endosoma" redirects here. For Endosomaphilia, see <a href="/wiki/Vorarephilia" title="Vorarephilia">Vorarephilia</a>.</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg" class="mw-file-description"><img alt="endocytic pathway compartments" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg/315px-HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg" decoding="async" width="315" height="339" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg/473px-HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/85/HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg/630px-HeLa_cell_endocytic_pathway_labeled_for_EGFR_and_transferrin.jpg 2x" data-file-width="3259" data-file-height="3510" /></a><figcaption>Electron micrograph of endosomes in human <a href="/wiki/HeLa_cell" class="mw-redirect" title="HeLa cell">HeLa cells</a>. Early endosomes (E - labeled for EGFR, 5 minutes after internalisation, and transferrin), late endosomes/MVBs (M) and lysosomes (L) are visible. Bar, 500 nm.</figcaption></figure> <p><b>Endosomes</b> are a collection of intracellular sorting <a href="/wiki/Organelles" class="mw-redirect" title="Organelles">organelles</a> in <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a>. They are parts of the <a href="/wiki/Endocytosis" title="Endocytosis">endocytic</a> membrane transport pathway originating from the <a href="/wiki/Trans_Golgi_network" class="mw-redirect" title="Trans Golgi network">trans Golgi network</a>. Molecules or ligands internalized from the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> can follow this pathway all the way to <a href="/wiki/Lysosome" title="Lysosome">lysosomes</a> for degradation or can be recycled back to the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> in the <a href="/wiki/Endocytic_cycle" class="mw-redirect" title="Endocytic cycle">endocytic cycle</a>. Molecules are also transported to endosomes from the trans Golgi network and either continue to lysosomes or recycle back to the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a>. </p><p>Endosomes can be classified as early, sorting, or late depending on their stage post internalization.<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> Endosomes represent a major sorting compartment of the <a href="/wiki/Endomembrane_system" title="Endomembrane system">endomembrane system</a> in cells.<sup id="cite_ref-endo1_2-0" class="reference"><a href="#cite_note-endo1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endosome&action=edit&section=1" title="Edit section: Function"><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">Further information: <a href="/wiki/Endocytic_cycle" class="mw-redirect" title="Endocytic cycle">Endocytic cycle</a></div> <p>Endosomes provide an environment for material to be sorted before it reaches the degradative lysosome.<sup id="cite_ref-endo1_2-1" class="reference"><a href="#cite_note-endo1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/Low-density_lipoprotein" title="Low-density lipoprotein">low-density lipoprotein</a> (LDL) is taken into the cell by binding to the <a href="/wiki/LDL_receptor" title="LDL receptor">LDL receptor</a> at the cell surface. Upon reaching early endosomes, the LDL dissociates from the receptor, and the receptor can be recycled to the cell surface. The LDL remains in the endosome and is delivered to lysosomes for processing. LDL dissociates because of the slightly acidified environment of the early endosome, generated by a vacuolar membrane proton pump <a href="/wiki/V-ATPase" title="V-ATPase">V-ATPase</a>. On the other hand, <a href="/wiki/Epidermal_growth_factor" title="Epidermal growth factor">epidermal growth factor</a> (EGF) and the EGF receptor have a pH-resistant bond that persists until it is delivered to lysosomes for their degradation. The <a href="/wiki/Mannose_6-phosphate_receptor" title="Mannose 6-phosphate receptor">mannose 6-phosphate receptor</a> carries <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligands</a> from the Golgi destined for the lysosome by a similar mechanism. </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=Endosome&action=edit&section=2" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are three different types of endosomes: <i>early endosomes</i>, <i>late endosomes</i>, and <i>recycling endosomes</i>.<sup id="cite_ref-endo1_2-2" class="reference"><a href="#cite_note-endo1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> They are distinguished by the time it takes for endocytosed material to reach them, and by markers such as <a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rabs</a>.<sup id="cite_ref-Stenmark-2009_3-0" class="reference"><a href="#cite_note-Stenmark-2009-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> They also have different morphology. Once <a href="/wiki/Endocytosis" title="Endocytosis">endocytic</a> vesicles have uncoated, they fuse with early endosomes. Early endosomes then <i>mature</i> into late endosomes before fusing with lysosomes.<sup id="cite_ref-Futter-1996_4-0" class="reference"><a href="#cite_note-Futter-1996-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-eele2_5-0" class="reference"><a href="#cite_note-eele2-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Early endosomes mature in several ways to form late endosomes. They become increasingly acidic mainly through the activity of the V-ATPase.<sup id="cite_ref-Lafourcade-2008_6-0" class="reference"><a href="#cite_note-Lafourcade-2008-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Many molecules that are recycled are removed by concentration in the tubular regions of early endosomes. Loss of these tubules to recycling pathways means that late endosomes mostly lack tubules. They also increase in size due to the homotypic fusion of early endosomes into larger vesicles.<sup id="cite_ref-Rink-2005_7-0" class="reference"><a href="#cite_note-Rink-2005-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Molecules are also sorted into smaller vesicles that bud from the perimeter membrane into the endosome lumen, forming <b>intraluminal vesicles</b> (ILVs); this leads to the multivesicular appearance of late endosomes and so they are also known as <b>multivesicular endosomes</b> or <b>multivesicular bodies</b> (MVBs). Removal of recycling molecules such as <a href="/wiki/Transferrin_receptor" title="Transferrin receptor">transferrin receptors</a> and mannose 6-phosphate receptors continues during this period, probably via budding of vesicles out of endosomes.<sup id="cite_ref-Futter-1996_4-1" class="reference"><a href="#cite_note-Futter-1996-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Finally, the endosomes lose <a href="/wiki/RAB5A" title="RAB5A">RAB5A</a> and acquire <a href="/wiki/RAB7A" title="RAB7A">RAB7A</a>, making them competent for fusion with lysosomes.<sup id="cite_ref-Rink-2005_7-1" class="reference"><a href="#cite_note-Rink-2005-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Fusion of late endosomes with lysosomes has been shown to result in the formation of a 'hybrid' compartment, with characteristics intermediate of the two source compartments.<sup id="cite_ref-endo-lysfus_8-0" class="reference"><a href="#cite_note-endo-lysfus-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> For example, lysosomes are more dense than late endosomes, and the hybrids have an intermediate density. Lysosomes reform by recondensation to their normal, higher density. However, before this happens, more late endosomes may fuse with the hybrid. </p><p>Some material recycles to the plasma membrane directly from early endosomes,<sup id="cite_ref-fastrecyc1_9-0" class="reference"><a href="#cite_note-fastrecyc1-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> but most traffics via recycling endosomes. </p> <ul><li><i>Early endosomes</i> consist of a dynamic tubular-vesicular network (vesicles up to 1 μm in diameter with connected tubules of approx. 50 nm diameter). Markers include <a href="/wiki/RAB5A" title="RAB5A">RAB5A</a> and RAB4, <a href="/wiki/Transferrin" title="Transferrin">Transferrin</a> and its <a href="/wiki/Transferrin_receptor" title="Transferrin receptor">receptor</a> and <a href="/wiki/EEA1" title="EEA1">EEA1</a>.</li> <li><i>Late endosomes</i>, also known as MVBs, are mainly spherical, lack tubules, and contain many close-packed intraluminal vesicles. Markers include RAB7, RAB9, and mannose 6-phosphate receptors.<sup id="cite_ref-Russell-2006_10-0" class="reference"><a href="#cite_note-Russell-2006-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> In addition to this, the late endosomal membrane (and consequently the lysosome) contains a peculiar and unique <a href="/wiki/Lipid" title="Lipid">lipid</a> named BMP or LBPA, which is not found in any other organelle membrane.<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><sup id="cite_ref-Hullin-Matsuda_313–324_12-0" class="reference"><a href="#cite_note-Hullin-Matsuda_313–324-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup></li> <li><i>Recycling endosomes</i> are concentrated at the microtubule organizing center and consist of a mainly tubular network. Marker; RAB11.<sup id="cite_ref-Ullrich-1996_13-0" class="reference"><a href="#cite_note-Ullrich-1996-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup></li></ul> <p>More subtypes exist in specialized cells such as polarized cells and <a href="/wiki/Macrophage" title="Macrophage">macrophages</a>. </p><p><a href="/wiki/Phagosomes" class="mw-redirect" title="Phagosomes">Phagosomes</a>, <a href="/wiki/Macropinosome" title="Macropinosome">macropinosomes</a> and <a href="/wiki/Autophagosome" title="Autophagosome">autophagosomes</a><sup id="cite_ref-Fader-2009_14-0" class="reference"><a href="#cite_note-Fader-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> mature in a manner similar to endosomes, and may require fusion with normal endosomes for their maturation. Some intracellular pathogens subvert this process, for example, by preventing RAB7 acquisition.<sup id="cite_ref-Körner-2006_15-0" class="reference"><a href="#cite_note-Körner-2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Late endosomes/MVBs are sometimes called <i>endocytic carrier vesicles</i>, but this term was used to describe vesicles that bud from early endosomes and fuse with late endosomes. However, several observations (described above) have now demonstrated that it is more likely that transport between these two compartments occurs by a maturation process, rather than vesicle transport. </p><p>Another unique identifying feature that differs between the various classes of endosomes is the lipid composition in their membranes. Phosphatidyl inositol phosphates (PIPs), one of the most important <a href="/wiki/Lipid" title="Lipid">lipid</a> signaling molecules, is found to differ as the endosomes mature from early to late. <a href="/wiki/Phosphatidylinositol_4,5-bisphosphate" title="Phosphatidylinositol 4,5-bisphosphate">PI(4,5)P<sub>2</sub></a> is present on <a href="/wiki/Cell_membrane" title="Cell membrane">plasma membranes</a>, <a href="/wiki/Phosphatidylinositol_3-phosphate" title="Phosphatidylinositol 3-phosphate">PI(3)P</a> on early endosomes, PI(3,5)P<sub>2</sub> on late endosomes and <a href="/wiki/Phosphatidylinositol_4-phosphate" title="Phosphatidylinositol 4-phosphate">PI(4)P</a> on the <a href="/wiki/Trans_Golgi_network" class="mw-redirect" title="Trans Golgi network">trans Golgi network</a>.<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> These lipids on the surface of the endosomes help in the specific recruitment of proteins from the cytosol, thus providing them an identity. The inter-conversion of these lipids is a result of the concerted action of phosphoinositide <a href="/wiki/Kinase" title="Kinase">kinases</a> and <a href="/wiki/Phosphatases" class="mw-redirect" title="Phosphatases">phosphatases</a> that are strategically localized<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> </p> <div class="mw-heading mw-heading2"><h2 id="Pathways">Pathways</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endosome&action=edit&section=3" title="Edit section: Pathways"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Endocytic_pathway_of_animal_cells_showing_EGF_receptors,_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg" class="mw-file-description"><img alt="animal cell endocytic pathway" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg/315px-Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg" decoding="async" width="315" height="276" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg/473px-Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg/630px-Endocytic_pathway_of_animal_cells_showing_EGF_receptors%2C_transferrin_receptors_and_mannose-6-phosphate_receptors.jpg 2x" data-file-width="2112" data-file-height="1849" /></a><figcaption>Diagram of the pathways that intersect endosomes in the endocytic pathway of animal cells. Examples of molecules that follow some of the pathways are shown, including receptors for EGF, transferrin, and lysosomal hydrolases. Recycling endosomes, and compartments and pathways found in more specialized cells, are not shown.</figcaption></figure> <p>There are three main compartments that have pathways that connect with endosomes. More pathways exist in specialized cells, such as <a href="/wiki/Melanocytes" class="mw-redirect" title="Melanocytes">melanocytes</a> and polarized cells. For example, in <a href="/wiki/Epithelium" title="Epithelium">epithelial</a> cells, a special process called <a href="/wiki/Transcytosis" title="Transcytosis">transcytosis</a> allows some materials to enter one side of a cell and exit from the opposite side. Also, in some circumstances, late endosomes/MVBs fuse with the plasma membrane instead of with lysosomes, releasing the lumenal vesicles, now called <a href="/wiki/Exosome_(vesicle)" title="Exosome (vesicle)">exosomes</a>, into the extracellular medium. </p><p>There is no consensus as to the exact nature of these pathways, and the sequential route taken by any given cargo in any given situation will tend to be a matter of debate. </p> <div class="mw-heading mw-heading3"><h3 id="Golgi_to/from_endosomes"><span id="Golgi_to.2Ffrom_endosomes"></span>Golgi to/from endosomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endosome&action=edit&section=4" title="Edit section: Golgi to/from endosomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Vesicles pass between the Golgi and endosomes in both directions. The <a href="/wiki/GGA1" title="GGA1">GGAs</a> and <a href="/wiki/AP1G1" title="AP1G1">AP-1</a> <a href="/wiki/Clathrin-coated_vesicle" class="mw-redirect" title="Clathrin-coated vesicle">clathrin-coated vesicle</a> adaptors make vesicles at the Golgi that carry molecules to endosomes.<sup id="cite_ref-Ghosh-2004_18-0" class="reference"><a href="#cite_note-Ghosh-2004-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> In the opposite direction, <a href="/wiki/Retromer" title="Retromer">retromer</a> generates vesicles at early endosomes that carry molecules back to the Golgi. Some studies describe a retrograde traffic pathway from late endosomes to the Golgi that is mediated by <a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rab9</a> and <a href="/wiki/TIP47" class="mw-redirect" title="TIP47">TIP47</a>, but other studies dispute these findings. Molecules that follow these pathways include the mannose-6-phosphate receptors that carry lysosomal hydrolases to the endocytic pathway. The hydrolases are released in the acidic environment of endosomes, and the receptor is retrieved to the Golgi by retromer and Rab9. </p> <div class="mw-heading mw-heading3"><h3 id="Plasma_membrane_to/from_early_endosomes_(via_recycling_endosomes)"><span id="Plasma_membrane_to.2Ffrom_early_endosomes_.28via_recycling_endosomes.29"></span>Plasma membrane to/from early endosomes (via recycling endosomes)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endosome&action=edit&section=5" title="Edit section: Plasma membrane to/from early endosomes (via recycling endosomes)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Molecules are delivered from the plasma membrane to early endosomes in <a href="/wiki/Endocytosis" title="Endocytosis">endocytic</a> vesicles. Molecules can be internalized via <a href="/wiki/Receptor-mediated_endocytosis" title="Receptor-mediated endocytosis">receptor-mediated endocytosis</a> in <a href="/wiki/Clathrin" title="Clathrin">clathrin</a>-coated vesicles. Other types of vesicles also form at the plasma membrane for this pathway, including ones utilising <a href="/wiki/Caveolae" title="Caveolae">caveolin</a>. Vesicles also transport molecules directly back to the plasma membrane, but many molecules are transported in vesicles that first fuse with recycling endosomes.<sup id="cite_ref-Grant-2009_19-0" class="reference"><a href="#cite_note-Grant-2009-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Molecules following this recycling pathway are concentrated in the tubules of early endosomes. Molecules that follow these pathways include the <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a> for <a href="/wiki/LDL" class="mw-redirect" title="LDL">LDL</a>, <a href="/wiki/Epidermal_growth_factor" title="Epidermal growth factor">epidermal growth factor</a> (EGF), and the iron transport protein transferrin. Internalization of these receptors from the plasma membrane occurs by receptor-mediated endocytosis. LDL is released in endosomes because of the lower pH, and the receptor is recycled to the cell surface. <a href="/wiki/Cholesterol" title="Cholesterol">Cholesterol</a> is carried in the blood primarily by (LDL), and transport by the LDL receptor is the main mechanism by which cholesterol is taken up by cells. EGFRs are activated when EGF binds. The activated receptors stimulate their own internalization and degradation in lysosomes. EGF remains bound to the <a href="/wiki/EGF_receptor" class="mw-redirect" title="EGF receptor">EGF receptor</a> (EGFR) once it is endocytosed to endosomes. The activated EGFRs stimulate their own ubiquitination, and this directs them to lumenal vesicles (see below) and so they are not recycled to the plasma membrane. This removes the signaling portion of the protein from the cytosol and thus prevents continued stimulation of growth<sup id="cite_ref-Futter-2001_20-0" class="reference"><a href="#cite_note-Futter-2001-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> - in cells not stimulated with EGF, EGFRs have no EGF bound to them and therefore recycle if they reach endosomes.<sup id="cite_ref-Felder-1990_21-0" class="reference"><a href="#cite_note-Felder-1990-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> Transferrin also remains associated with its receptor, but, in the acidic endosome, iron is released from the transferrin, and then the iron-free transferrin (still bound to the transferrin receptor) returns from the early endosome to the cell surface, both directly and via recycling endosomes.<sup id="cite_ref-Dautry-Varsat-1986_22-0" class="reference"><a href="#cite_note-Dautry-Varsat-1986-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Late_endosomes_to_lysosomes">Late endosomes to lysosomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endosome&action=edit&section=6" title="Edit section: Late endosomes to lysosomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Transport from late endosomes to lysosomes is, in essence, unidirectional, since a late endosome is "consumed" in the process of fusing with a lysosome (sometimes called endolysosome<sup id="cite_ref-pmid34611326_23-0" class="reference"><a href="#cite_note-pmid34611326-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid33324224_24-0" class="reference"><a href="#cite_note-pmid33324224-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup>).Hence, soluble molecules in the lumen of endosomes will tend to end up in lysosomes, unless they are retrieved in some way. <a href="/wiki/Transmembrane_protein" title="Transmembrane protein">Transmembrane proteins</a> can be delivered to the perimeter membrane or the lumen of lysosomes. Transmembrane proteins destined for the lysosome lumen are sorted into the vesicles that bud from the perimeter membrane into endosomes, a process that begins in early endosomes. The process of creating vesicles within the endosome is thought to be enhanced by the peculiar lipid BMP or LBPA, which is only found in late endosomes, endolysosomes or lysosomes.<sup id="cite_ref-Hullin-Matsuda_313–324_12-1" class="reference"><a href="#cite_note-Hullin-Matsuda_313–324-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> When the endosome has matured into a late endosome/MVB and fuses with a lysosome, the vesicles in the lumen are delivered to the lysosome lumen. Proteins are marked for this pathway by the addition of <a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitin</a>.<sup id="cite_ref-Hicke-2003_25-0" class="reference"><a href="#cite_note-Hicke-2003-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Endosomal_sorting_complexes_required_for_transport" class="mw-redirect" title="Endosomal sorting complexes required for transport">endosomal sorting complexes required for transport</a> (ESCRTs) recognise this ubiquitin and sort the protein into the forming lumenal vesicles.<sup id="cite_ref-Hurley-2008_26-0" class="reference"><a href="#cite_note-Hurley-2008-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Molecules that follow these pathways include LDL and the lysosomal hydrolases delivered by mannose-6-phosphate receptors. These soluble molecules remain in endosomes and are therefore delivered to lysosomes. Also, the transmembrane EGFRs, bound to EGF, are tagged with ubiquitin and are therefore sorted into lumenal vesicles by the ESCRTs. </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=Endosome&action=edit&section=7" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Back-Fusion" title="Back-Fusion">Back-Fusion</a></li> <li><a href="/wiki/Exosome_(vesicle)" title="Exosome (vesicle)">Exosome</a></li> <li><a href="/wiki/Paramural_body" title="Paramural body">Paramural body</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=Endosome&action=edit&section=8" 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-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="CITEREFStoorvogelStrousGeuzeOorschot1991" class="citation journal cs1">Stoorvogel W, Strous GJ, Geuze HJ, Oorschot V, Schwartz AL (May 1991). 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"Regulation of membrane protein transport by ubiquitin and ubiquitin-binding proteins". <i>Annual Review of Cell and Developmental Biology</i>. <b>19</b>: <span class="nowrap">141–</span>72. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.cellbio.19.110701.154617">10.1146/annurev.cellbio.19.110701.154617</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/14570567">14570567</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+Cell+and+Developmental+Biology&rft.atitle=Regulation+of+membrane+protein+transport+by+ubiquitin+and+ubiquitin-binding+proteins&rft.volume=19&rft.pages=%3Cspan+class%3D%22nowrap%22%3E141-%3C%2Fspan%3E72&rft.date=2003&rft_id=info%3Adoi%2F10.1146%2Fannurev.cellbio.19.110701.154617&rft_id=info%3Apmid%2F14570567&rft.aulast=Hicke&rft.aufirst=L&rft.au=Dunn%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndosome" class="Z3988"></span></span> </li> <li id="cite_note-Hurley-2008-26"><span class="mw-cite-backlink"><b><a href="#cite_ref-Hurley-2008_26-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHurley2008" class="citation journal cs1">Hurley JH (February 2008). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2282067">"ESCRT complexes and the biogenesis of multivesicular bodies"</a>. <i>Current Opinion in Cell Biology</i>. <b>20</b> (1): <span class="nowrap">4–</span>11. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ceb.2007.12.002">10.1016/j.ceb.2007.12.002</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/PMC2282067">2282067</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/18222686">18222686</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Opinion+in+Cell+Biology&rft.atitle=ESCRT+complexes+and+the+biogenesis+of+multivesicular+bodies&rft.volume=20&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E4-%3C%2Fspan%3E11&rft.date=2008-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2282067%23id-name%3DPMC&rft_id=info%3Apmid%2F18222686&rft_id=info%3Adoi%2F10.1016%2Fj.ceb.2007.12.002&rft.aulast=Hurley&rft.aufirst=JH&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2282067&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndosome" class="Z3988"></span></span> </li> </ol></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlberts2004" class="citation book cs1">Alberts, Bruce; et al. (2004). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/essentialcellbio00albe"><i>Essential Cell Biology</i></a></span> (2nd ed.). New York, NY: Garland Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-3480-4" title="Special:BookSources/978-0-8153-3480-4"><bdi>978-0-8153-3480-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Essential+Cell+Biology&rft.place=New+York%2C+NY&rft.edition=2nd&rft.pub=Garland+Science&rft.date=2004&rft.isbn=978-0-8153-3480-4&rft.aulast=Alberts&rft.aufirst=Bruce&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fessentialcellbio00albe&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndosome" class="Z3988"></span></li></ul> <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=Endosome&action=edit&section=9" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://opm.phar.umich.edu/localization.php?localization=Endosome%20membrane">3D structures of some proteins associated with endosome membrane</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul 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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 class="mw-selflink selflink">Endosome</a></li> <li><a href="/wiki/Phagosome" title="Phagosome">Phagosome</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Acrosome" title="Acrosome">Acrosome</a></li></ul></li> <li><a href="/wiki/Granule_(cell_biology)" title="Granule (cell biology)">Cytoplasmic granule</a> <ul><li><a href="/wiki/Melanosome" title="Melanosome">Melanosome</a></li> <li><a href="/wiki/Microbody" title="Microbody">Microbody</a></li> <li><a href="/wiki/Glyoxysome" title="Glyoxysome">Glyoxysome</a></li> <li><a href="/wiki/Peroxisome" title="Peroxisome">Peroxisome</a></li> <li><a href="/wiki/Weibel%E2%80%93Palade_body" title="Weibel–Palade body">Weibel–Palade body</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Microfilament" title="Microfilament">Microfilament</a></li> <li><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate filament</a></li> <li><a href="/wiki/Microtubule" title="Microtubule">Microtubule</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a></li> <li><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organizing center</a> <ul><li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Centriole" title="Centriole">Centriole</a></li> <li><a href="/wiki/Basal_body" title="Basal body">Basal body</a></li> <li><a href="/wiki/Spindle_pole_body" title="Spindle pole body">Spindle pole body</a></li></ul></li> <li><a href="/wiki/Myofibril" title="Myofibril">Myofibril</a></li> <li><a href="/wiki/Undulipodium" title="Undulipodium">Undulipodium</a> <ul><li><a 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><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?useformat=desktop&type=1x1&usesul3=0" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Endosome&oldid=1226214717">https://en.wikipedia.org/w/index.php?title=Endosome&oldid=1226214717</a>"</div></div> <div id="catlinks" class="catlinks" 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