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id="toc-Anaphase_A" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anaphase_A"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Anaphase A</span> </div> </a> <ul id="toc-Anaphase_A-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Anaphase_B" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anaphase_B"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Anaphase B</span> </div> </a> <ul id="toc-Anaphase_B-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Relation_to_the_cell_cycle" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Relation_to_the_cell_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Relation to the cell cycle</span> </div> </a> <ul id="toc-Relation_to_the_cell_cycle-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">3</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">4</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">5</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> 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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">Anaphase</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 33 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-33" 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">33 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%B7%D9%88%D8%B1_%D8%A7%D9%84%D8%B5%D8%B9%D9%88%D8%AF" title="طور الصعود – Arabic" lang="ar" hreflang="ar" data-title="طور الصعود" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Anafaza" title="Anafaza – Azerbaijani" lang="az" hreflang="az" data-title="Anafaza" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%85%E0%A7%8D%E0%A6%AF%E0%A6%BE%E0%A6%A8%E0%A6%BE%E0%A6%AB%E0%A7%87%E0%A6%9C" title="অ্যানাফেজ – Bangla" lang="bn" hreflang="bn" data-title="অ্যানাফেজ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%90%D0%BD%D0%B0%D1%84%D0%B0%D0%B7%D0%B0" title="Анафаза – Bulgarian" lang="bg" hreflang="bg" data-title="Анафаза" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Anafaza_(mitoza)" title="Anafaza (mitoza) – Bosnian" lang="bs" hreflang="bs" data-title="Anafaza (mitoza)" 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/Anafase" title="Anafase – Catalan" lang="ca" hreflang="ca" data-title="Anafase" 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/Anaf%C3%A1ze" title="Anafáze – Czech" lang="cs" hreflang="cs" data-title="Anafáze" 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/Mitose#Anaphase" title="Mitose – German" lang="de" hreflang="de" data-title="Mitose" 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%91%CE%BD%CE%AC%CF%86%CE%B1%CF%83%CE%B7" title="Ανάφαση – Greek" lang="el" hreflang="el" data-title="Ανάφαση" 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/Anafase" title="Anafase – Spanish" lang="es" hreflang="es" data-title="Anafase" 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%A2%D9%86%D8%A7%D9%81%D8%A7%D8%B2" title="آنافاز – Persian" lang="fa" hreflang="fa" data-title="آنافاز" 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/Anaphase" title="Anaphase – French" lang="fr" hreflang="fr" data-title="Anaphase" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B1%D5%B6%D5%A1%D6%86%D5%A1%D5%A6" title="Անաֆազ – Armenian" lang="hy" hreflang="hy" data-title="Անաֆազ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Anafaza" title="Anafaza – Croatian" lang="hr" hreflang="hr" data-title="Anafaza" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Anafazo" title="Anafazo – Ido" lang="io" hreflang="io" data-title="Anafazo" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%85%E0%B2%A8%E0%B2%BE%E0%B2%AB%E0%B3%87%E0%B2%B8%E0%B3%8D" title="ಅನಾಫೇಸ್ – Kannada" lang="kn" hreflang="kn" data-title="ಅನಾಫೇಸ್" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%90%E1%83%9C%E1%83%90%E1%83%A4%E1%83%90%E1%83%96%E1%83%90" title="ანაფაზა – Georgian" lang="ka" hreflang="ka" data-title="ანაფაზა" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%90%D0%BD%D0%B0%D1%84%D0%B0%D0%B7%D0%B0" title="Анафаза – Kazakh" lang="kk" hreflang="kk" data-title="Анафаза" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Anafaz" title="Anafaz – Haitian Creole" lang="ht" hreflang="ht" data-title="Anafaz" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Anafasa" title="Anafasa – Malay" lang="ms" hreflang="ms" data-title="Anafasa" 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/Anafase" title="Anafase – Dutch" lang="nl" hreflang="nl" data-title="Anafase" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%BE%8C%E6%9C%9F_(%E7%B4%B0%E8%83%9E%E5%88%86%E8%A3%82)" title="後期 (細胞分裂) – Japanese" lang="ja" hreflang="ja" data-title="後期 (細胞分裂)" 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-om mw-list-item"><a href="https://om.wikipedia.org/wiki/Anaafeezii" title="Anaafeezii – Oromo" lang="om" hreflang="om" data-title="Anaafeezii" data-language-autonym="Oromoo" data-language-local-name="Oromo" class="interlanguage-link-target"><span>Oromoo</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Anafaza" title="Anafaza – Uzbek" lang="uz" hreflang="uz" data-title="Anafaza" 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/Anafaza" title="Anafaza – Polish" lang="pl" hreflang="pl" data-title="Anafaza" 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/An%C3%A1fase" title="Anáfase – Portuguese" lang="pt" hreflang="pt" data-title="Anáfase" 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/Anafaz%C4%83" title="Anafază – Romanian" lang="ro" hreflang="ro" data-title="Anafază" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17559452 badge-recommendedarticle mw-list-item" title="recommended article"><a href="https://ru.wikipedia.org/wiki/%D0%90%D0%BD%D0%B0%D1%84%D0%B0%D0%B7%D0%B0" title="Анафаза – Russian" lang="ru" hreflang="ru" data-title="Анафаза" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Anafaza" title="Anafaza – Serbian" lang="sr" hreflang="sr" data-title="Anafaza" 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-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Anapase" title="Anapase – Tagalog" lang="tl" hreflang="tl" data-title="Anapase" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Anafaz" title="Anafaz – Turkish" lang="tr" hreflang="tr" data-title="Anafaz" 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%90%D0%BD%D0%B0%D1%84%D0%B0%D0%B7%D0%B0" title="Анафаза – Ukrainian" lang="uk" hreflang="uk" data-title="Анафаза" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/K%E1%BB%B3_sau" title="Kỳ sau – Vietnamese" lang="vi" hreflang="vi" data-title="Kỳ sau" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</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/Q22299157#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> 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.hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Anaphase_I" class="mw-redirect" title="Anaphase I">Anaphase I</a> or <a href="/wiki/Anaphase_II" class="mw-redirect" title="Anaphase II">Anaphase II</a>.</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Anaphase.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Anaphase.svg/330px-Anaphase.svg.png" decoding="async" width="300" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Anaphase.svg/500px-Anaphase.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/Anaphase.svg/960px-Anaphase.svg.png 2x" data-file-width="2680" data-file-height="1400" /></a><figcaption></figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Anaphase_IF.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Anaphase_IF.jpg/330px-Anaphase_IF.jpg" decoding="async" width="300" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Anaphase_IF.jpg/450px-Anaphase_IF.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/9/91/Anaphase_IF.jpg 2x" data-file-width="480" data-file-height="480" /></a><figcaption>A cell during anaphase. Microtubules are visible in green.</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Stages_of_late_M_phase_in_a_vertebrate_cell.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Stages_of_late_M_phase_in_a_vertebrate_cell.svg/250px-Stages_of_late_M_phase_in_a_vertebrate_cell.svg.png" decoding="async" width="220" height="684" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Stages_of_late_M_phase_in_a_vertebrate_cell.svg/330px-Stages_of_late_M_phase_in_a_vertebrate_cell.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Stages_of_late_M_phase_in_a_vertebrate_cell.svg/500px-Stages_of_late_M_phase_in_a_vertebrate_cell.svg.png 2x" data-file-width="512" data-file-height="1593" /></a><figcaption>Stages of late M phase in a vertebrate cell</figcaption></figure> <p><b>Anaphase</b> (from <a href="/wiki/Ancient_Greek_language" class="mw-redirect" title="Ancient Greek language">Ancient Greek</a> <i> </i><a href="https://en.wiktionary.org/wiki/%E1%BC%80%CE%BD%CE%B1-" class="extiw" title="wikt:ἀνα-">ἀνα-</a><i> (<a href="https://en.wiktionary.org/wiki/ana-" class="extiw" title="wikt:ana-">ana-</a>)</i> <span class="gloss-quot">'</span><span class="gloss-text">back, backward</span><span class="gloss-quot">'</span> and <i> </i><a href="https://en.wiktionary.org/wiki/%CF%86%CE%AC%CF%83%CE%B9%CF%82" class="extiw" title="wikt:φάσις">φάσις</a><i> (phásis)</i> <span class="gloss-quot">'</span><span class="gloss-text">appearance</span><span class="gloss-quot">'</span>) is the stage of <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> after the process of <a href="/wiki/Metaphase" title="Metaphase">metaphase</a>, when replicated <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> are split and the newly-copied chromosomes (daughter <a href="/wiki/Chromatid" title="Chromatid">chromatids</a>) are moved to opposite poles of the cell. <a href="/wiki/Chromosome" title="Chromosome">Chromosomes</a> also reach their overall maximum condensation in late anaphase, to help <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> segregation and the re-formation of the nucleus.<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> </p><p>Anaphase starts when the <a href="/wiki/Anaphase_promoting_complex" class="mw-redirect" title="Anaphase promoting complex">anaphase promoting complex</a> marks an <a href="/wiki/Chaperone_(protein)" title="Chaperone (protein)">inhibitory chaperone</a> called <a href="/wiki/Securin" title="Securin">securin</a> for destruction by <a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitylating</a> it. Securin is a protein which inhibits a <a href="/wiki/Protease" title="Protease">protease</a> known as <a href="/wiki/Separase" title="Separase">separase</a>. The destruction of securin unleashes separase which then breaks down <a href="/wiki/Cohesin" title="Cohesin">cohesin</a>, a protein responsible for holding sister chromatids together.<sup id="cite_ref-Kimball's_Biology_Pages_2-0" class="reference"><a href="#cite_note-Kimball's_Biology_Pages-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>At this point, three subclasses of <a href="/wiki/Microtubule" title="Microtubule">microtubule</a> unique to mitosis are involved in creating the forces necessary to separate the chromatids: kinetochore microtubules, interpolar microtubules, and <a href="/wiki/Aster_(cell_biology)#Astral_microtubules" title="Aster (cell biology)">astral</a> microtubules. </p><p>The centromeres are split, and the sister chromatids are pulled toward the poles by kinetochore microtubules. They take on a V-shape or Y-shape as they are pulled to either pole. </p><p>While the chromosomes are drawn to each side of the cell, interpolar microtubules and astral microtubules generate forces that stretch the cell into an oval.<sup id="cite_ref-NCBI_3-0" class="reference"><a href="#cite_note-NCBI-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Once anaphase is complete, the cell moves into <a href="/wiki/Telophase" title="Telophase">telophase</a>.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Phases">Phases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Anaphase&action=edit&section=1" title="Edit section: Phases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Anaphase is characterized by two distinct motions. The first of these, anaphase A, moves chromosomes to either pole of a dividing cell (marked by <a href="/wiki/Centrosome" title="Centrosome">centrosomes</a>, from which mitotic microtubules are generated and organised). The movement for this is primarily generated by the action of kinetochores, and a subclass of microtubule called kinetochore microtubules. </p><p>The second motion, anaphase B, involves the separation of these poles from each other. The movement for this is primarily generated by the action of interpolar microtubules and astral microtubules. </p> <div class="mw-heading mw-heading3"><h3 id="Anaphase_A">Anaphase A</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Anaphase&action=edit&section=2" title="Edit section: Anaphase A"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A combination of different forces have been observed acting on chromatids in anaphase A, but the primary force is exerted centrally. Microtubules attach to the midpoint of chromosomes (the <a href="/wiki/Centromere" title="Centromere">centromere</a>) via protein complexes (<a href="/wiki/Kinetochore" title="Kinetochore">kinetochores</a>). The attached microtubules depolymerise and shorten, which together with motor proteins creates movement that pulls chromosomes towards centrosomes located at each pole of the cell.<sup id="cite_ref-:0_5-0" class="reference"><a href="#cite_note-:0-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Anaphase_B">Anaphase B</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Anaphase&action=edit&section=3" title="Edit section: Anaphase B"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The second part of anaphase is driven by its own distinct mechanisms. Force is generated by several actions. Interpolar microtubules begin at each centrosome and join at the equator of the dividing cell. They push against one another, causing each centrosome to move further apart. Meanwhile, astral microtubules begin at each centrosome and join with the cell membrane. This allows them to pull each centrosome closer to the cell membrane. Movement created by these microtubules is generated by a combination of microtubule growth or shrinking, and by motor proteins such as <a href="/wiki/Dynein" title="Dynein">dyneins</a> or <a href="/wiki/Kinesin" title="Kinesin">kinesins</a>.<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> <div class="mw-heading mw-heading2"><h2 id="Relation_to_the_cell_cycle">Relation to the cell cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Anaphase&action=edit&section=4" title="Edit section: Relation to the cell cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Anaphase accounts for approximately 1% of the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>'s duration.<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> It begins with the regulated triggering of the metaphase-to-anaphase transition. <a href="/wiki/Metaphase" title="Metaphase">Metaphase</a> ends with the destruction of B <a href="/wiki/Cyclin" title="Cyclin">cyclin</a>. B cyclin is marked with <a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitin</a> which flags it for destruction by <a href="/wiki/Proteasomes" class="mw-redirect" title="Proteasomes">proteasomes</a>, which is required for the function of metaphase cyclin-dependent kinases (M-Cdks). In essence, Activation of the <a href="/wiki/Anaphase-promoting_complex" title="Anaphase-promoting complex">Anaphase-promoting complex</a> (APC) causes the APC to cleave the M-phase cyclin and the inhibitory protein <a href="/wiki/Securin" title="Securin">securin</a> which activates the separase protease to cleave the <a href="/wiki/Cohesin" title="Cohesin">cohesin</a> subunits holding the <a href="/wiki/Chromatids" class="mw-redirect" title="Chromatids">chromatids</a> together. </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=Anaphase&action=edit&section=5" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Interphase" title="Interphase">Interphase</a></li> <li><a href="/wiki/Prophase" title="Prophase">Prophase</a></li> <li><a href="/wiki/Prometaphase" title="Prometaphase">Prometaphase</a></li> <li><a href="/wiki/Metaphase" title="Metaphase">Metaphase</a></li> <li><a href="/wiki/Telophase" title="Telophase">Telophase</a></li> <li><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a></li> <li><a href="/wiki/Meiosis#Anaphase_I" title="Meiosis">Anaphase I</a></li> <li><a href="/wiki/Meiosis#Meiosis_II" title="Meiosis">Anaphase II</a></li> <li><a href="/wiki/Cdc20" class="mw-redirect" title="Cdc20">Cdc20</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=Anaphase&action=edit&section=6" 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 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Veterinary+Pathology&rft.atitle=The+cell+cycle%3A+a+review&rft.volume=35&rft.issue=6&rft.pages=%3Cspan+class%3D%22nowrap%22%3E461-%3C%2Fspan%3E78&rft.date=1998-11&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A43902779%23id-name%3DS2CID&rft_id=info%3Apmid%2F9823588&rft_id=info%3Adoi%2F10.1177%2F030098589803500601&rft.aulast=Schafer&rft.aufirst=KA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAnaphase" class="Z3988"></span></span> </li> <li id="cite_note-:0-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-:0_5-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAsbury2017" class="citation journal cs1">Asbury CL (February 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5372008">"Anaphase A: Disassembling Microtubules Move 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biology&rft.atitle=Anaphase+A%3A+Disassembling+Microtubules+Move+Chromosomes+toward+Spindle+Poles&rft.volume=6&rft.issue=1&rft.pages=15&rft.date=2017-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5372008%23id-name%3DPMC&rft_id=info%3Apmid%2F28218660&rft_id=info%3Adoi%2F10.3390%2Fbiology6010015&rft.aulast=Asbury&rft.aufirst=CL&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5372008&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAnaphase" 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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFScholeyCivelekoglu-ScholeyBrust-Mascher2016" class="citation journal cs1">Scholey JM, Civelekoglu-Scholey G, Brust-Mascher I (December 2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5192431">"Anaphase B"</a>. <i>Biology</i>. <b>5</b> (4): 51. <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%2Fbiology5040051">10.3390/biology5040051</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/PMC5192431">5192431</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/27941648">27941648</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biology&rft.atitle=Anaphase+B&rft.volume=5&rft.issue=4&rft.pages=51&rft.date=2016-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5192431%23id-name%3DPMC&rft_id=info%3Apmid%2F27941648&rft_id=info%3Adoi%2F10.3390%2Fbiology5040051&rft.aulast=Scholey&rft.aufirst=JM&rft.au=Civelekoglu-Scholey%2C+G&rft.au=Brust-Mascher%2C+I&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5192431&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAnaphase" class="Z3988"></span></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFHeathRethoret1980" class="citation journal cs1">Heath IB, Rethoret K (June 1980). "Temporal analysis of the nuclear cycle by serial section electron microscopy of the fungus, Saprolegnia ferax". <i>European Journal of Cell Biology</i>. <b>21</b> (2): <span class="nowrap">208–</span>13. <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/7398661">7398661</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=European+Journal+of+Cell+Biology&rft.atitle=Temporal+analysis+of+the+nuclear+cycle+by+serial+section+electron+microscopy+of+the+fungus%2C+Saprolegnia+ferax&rft.volume=21&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E208-%3C%2Fspan%3E13&rft.date=1980-06&rft_id=info%3Apmid%2F7398661&rft.aulast=Heath&rft.aufirst=IB&rft.au=Rethoret%2C+K&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAnaphase" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Anaphase&action=edit&section=7" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Commons-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/20px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/40px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span> Media related to <a href="https://commons.wikimedia.org/wiki/Category:Anaphase" class="extiw" title="commons:Category:Anaphase">Anaphase</a> at Wikimedia Commons</li></ul> <div class="navbox-styles"><style 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title="Protein">proteins</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cyclin" title="Cyclin">Cyclin</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/Cyclin_A" title="Cyclin A">A</a> (<a href="/wiki/Cyclin_A1" title="Cyclin A1">A1</a>, <a href="/wiki/Cyclin_A2" title="Cyclin A2">A2</a>)</li> <li><a href="/wiki/Cyclin_B" title="Cyclin B">B</a> (<a href="/wiki/Cyclin_B1" title="Cyclin B1">B1</a>, <a href="/wiki/Cyclin_B2" title="Cyclin B2">B2</a>, B3)</li> <li><a href="/wiki/Cyclin_D" title="Cyclin D">D</a> (<a href="/wiki/Cyclin_D1" title="Cyclin D1">D1</a>, <a href="/wiki/Cyclin_D2" title="Cyclin D2">D2</a>, <a href="/wiki/Cyclin_D3" title="Cyclin D3">D3</a>)</li> <li><a href="/wiki/Cyclin_E" title="Cyclin E">E</a> (<a href="/wiki/Cyclin_E1" title="Cyclin E1">E1</a>, <a href="/wiki/Cyclin_E2" title="Cyclin E2">E2</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">CDK</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/Cyclin-dependent_kinase_1" title="Cyclin-dependent kinase 1">1</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_2" title="Cyclin-dependent kinase 2">2</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_3" title="Cyclin-dependent kinase 3">3</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">4</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_5" title="Cyclin-dependent kinase 5">5</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_6" title="Cyclin-dependent kinase 6">6</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_7" title="Cyclin-dependent kinase 7">7</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_8" title="Cyclin-dependent kinase 8">8</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_9" title="Cyclin-dependent kinase 9">9</a></li> <li><a href="/wiki/Cyclin-dependent_kinase_10" title="Cyclin-dependent kinase 10">10</a></li> <li><a href="/wiki/CDC2L2" title="CDC2L2">11A</a></li> <li><a href="/wiki/CDC2L1" title="CDC2L1">11B</a></li> <li><a href="/wiki/CDK12" title="CDK12">12</a></li> <li><a href="/wiki/CDK13" title="CDK13">13</a></li> <li><a href="/wiki/PFTK1" title="PFTK1">14</a></li> <li><a href="/wiki/CDK-activating_kinase" title="CDK-activating kinase">CDK-activating kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cyclin-dependent_kinase_inhibitor_protein" title="Cyclin-dependent kinase inhibitor protein">CDK inhibitor</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/INK4" title="INK4">INK4a/ARF</a> (<a href="/wiki/P14arf" title="P14arf">p14arf</a>/<a href="/wiki/P16" title="P16">p16</a>, <a href="/wiki/CDKN2B" title="CDKN2B">p15</a>, <a href="/wiki/CDKN2C" title="CDKN2C">p18</a>, <a href="/wiki/CDKN2D" title="CDKN2D">p19</a>)</li> <li><a href="/wiki/Cell_cycle#Inhibitors" title="Cell cycle">cip/kip</a> (<a href="/wiki/P21" title="P21">p21</a>, <a href="/wiki/CDKN1B" title="CDKN1B">p27</a>, <a href="/wiki/Cyclin-dependent_kinase_inhibitor_1C" title="Cyclin-dependent kinase inhibitor 1C">p57</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/P53_p63_p73_family" title="P53 p63 p73 family">P53 p63 p73 family</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/P53" title="P53">p53</a></li> <li><a href="/wiki/TP63" title="TP63">p63</a></li> <li><a href="/wiki/P73" title="P73">p73</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-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cdk1" class="mw-redirect" title="Cdk1">Cdc2</a></li> <li><a href="/wiki/Cdc25" title="Cdc25">Cdc25</a></li> <li><a href="/wiki/CDC42" title="CDC42">Cdc42</a></li> <li><a href="/wiki/Cellular_apoptosis_susceptibility_protein" title="Cellular apoptosis susceptibility protein">Cellular apoptosis susceptibility protein</a></li> <li><a href="/wiki/E2F" title="E2F">E2F</a></li> <li><a href="/wiki/Maturation_promoting_factor" title="Maturation promoting factor">Maturation promoting factor</a></li> <li><a href="/wiki/Wee1" title="Wee1">Wee</a></li> <li><a href="/wiki/Cullin" title="Cullin">Cullin</a> (<a href="/wiki/CUL7" title="CUL7">CUL7</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Phases and<br />checkpoints</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Interphase" title="Interphase">Interphase</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/G1_phase" title="G1 phase">G<sub>1</sub> phase</a></li> <li><a href="/wiki/S_phase" title="S phase">S phase</a></li> <li><a href="/wiki/G2_phase" title="G2 phase">G<sub>2</sub> phase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_division" title="Cell division">M phase</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/Mitosis" title="Mitosis">Mitosis</a> (<a href="/wiki/Preprophase" title="Preprophase">Preprophase</a></li> <li><a href="/wiki/Prophase" title="Prophase">Prophase</a></li> <li><a href="/wiki/Prometaphase" title="Prometaphase">Prometaphase</a></li> <li><a href="/wiki/Metaphase" title="Metaphase">Metaphase</a></li> <li><a class="mw-selflink selflink">Anaphase</a></li> <li><a href="/wiki/Telophase" title="Telophase">Telophase</a>)</li> <li><a href="/wiki/Cytokinesis" title="Cytokinesis">Cytokinesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">Cell cycle checkpoints</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/Restriction_point" title="Restriction point">Restriction point</a></li> <li><a href="/wiki/Spindle_checkpoint" title="Spindle checkpoint">Spindle checkpoint</a></li> <li><a href="/wiki/Postreplication_checkpoint" title="Postreplication checkpoint">Postreplication checkpoint</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other cellular phases</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/Apoptosis" title="Apoptosis">Apoptosis</a></li> <li><a href="/wiki/G0_phase" title="G0 phase">G<sub>0</sub> phase</a></li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a></li></ul> </div></td></tr></tbody></table><div></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=Anaphase&oldid=1221803917">https://en.wikipedia.org/w/index.php?title=Anaphase&oldid=1221803917</a>"</div></div> <div id="catlinks" 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