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class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#G0_phase_(quiescence)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>G<sub>0</sub> phase (quiescence)</span> </div> </a> <ul id="toc-G0_phase_(quiescence)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Interphase" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Interphase"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Interphase</span> </div> </a> <ul id="toc-Interphase-sublist" class="vector-toc-list"> <li id="toc-G1_phase_(First_growth_phase_or_Post_mitotic_gap_phase)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#G1_phase_(First_growth_phase_or_Post_mitotic_gap_phase)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2.1</span> <span>G<sub>1</sub> phase (First growth phase or Post mitotic gap phase)</span> </div> </a> <ul id="toc-G1_phase_(First_growth_phase_or_Post_mitotic_gap_phase)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-S_phase_(DNA_replication)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#S_phase_(DNA_replication)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2.2</span> <span>S phase (DNA replication)</span> </div> </a> <ul id="toc-S_phase_(DNA_replication)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-G2_phase_(growth)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#G2_phase_(growth)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2.3</span> <span>G<sub>2</sub> phase (growth)</span> </div> </a> <ul id="toc-G2_phase_(growth)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mitotic_phase_(chromosome_separation)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitotic_phase_(chromosome_separation)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Mitotic phase (chromosome separation)</span> </div> </a> <ul id="toc-Mitotic_phase_(chromosome_separation)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cytokinesis_phase_(separation_of_all_cell_components)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cytokinesis_phase_(separation_of_all_cell_components)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Cytokinesis phase (separation of all cell components)</span> </div> </a> <ul id="toc-Cytokinesis_phase_(separation_of_all_cell_components)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Regulation_of_eukaryotic_cell_cycle" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Regulation_of_eukaryotic_cell_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Regulation of eukaryotic cell cycle</span> </div> </a> <button aria-controls="toc-Regulation_of_eukaryotic_cell_cycle-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Regulation of eukaryotic cell cycle subsection</span> </button> <ul id="toc-Regulation_of_eukaryotic_cell_cycle-sublist" class="vector-toc-list"> <li id="toc-Role_of_cyclins_and_CDKs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_of_cyclins_and_CDKs"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Role of cyclins and CDKs</span> </div> </a> <ul id="toc-Role_of_cyclins_and_CDKs-sublist" class="vector-toc-list"> <li id="toc-General_mechanism_of_cyclin-CDK_interaction" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#General_mechanism_of_cyclin-CDK_interaction"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>General mechanism of cyclin-CDK interaction</span> </div> </a> <ul id="toc-General_mechanism_of_cyclin-CDK_interaction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Specific_action_of_cyclin-CDK_complexes" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Specific_action_of_cyclin-CDK_complexes"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Specific action of cyclin-CDK complexes</span> </div> </a> <ul id="toc-Specific_action_of_cyclin-CDK_complexes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Inhibitors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inhibitors"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Inhibitors</span> </div> </a> <ul id="toc-Inhibitors-sublist" class="vector-toc-list"> <li id="toc-Endogenous" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Endogenous"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.1</span> <span>Endogenous</span> </div> </a> <ul id="toc-Endogenous-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Synthetic" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Synthetic"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.2</span> <span>Synthetic</span> </div> </a> <ul id="toc-Synthetic-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Transcriptional_regulatory_network" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transcriptional_regulatory_network"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Transcriptional regulatory network</span> </div> </a> <ul id="toc-Transcriptional_regulatory_network-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_replication_and_DNA_replication_origin_activity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_replication_and_DNA_replication_origin_activity"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>DNA replication and DNA replication origin activity</span> </div> </a> <ul id="toc-DNA_replication_and_DNA_replication_origin_activity-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Checkpoints" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Checkpoints"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Checkpoints</span> </div> </a> <ul id="toc-Checkpoints-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Fluorescence_imaging_of_the_cell_cycle" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Fluorescence_imaging_of_the_cell_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Fluorescence imaging of the cell cycle</span> </div> </a> <ul id="toc-Fluorescence_imaging_of_the_cell_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_tumor_formation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_tumor_formation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Role in tumor formation</span> </div> </a> <ul id="toc-Role_in_tumor_formation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cell_cycle_evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Cell_cycle_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Cell cycle evolution</span> </div> </a> <button aria-controls="toc-Cell_cycle_evolution-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Cell cycle evolution subsection</span> </button> <ul id="toc-Cell_cycle_evolution-sublist" class="vector-toc-list"> <li id="toc-Evolution_of_the_genome" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution_of_the_genome"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Evolution of the genome</span> </div> </a> <ul id="toc-Evolution_of_the_genome-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cyclin-dependent_kinase_and_cyclin_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cyclin-dependent_kinase_and_cyclin_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Cyclin-dependent kinase and cyclin evolution</span> </div> </a> <ul id="toc-Cyclin-dependent_kinase_and_cyclin_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-G1/S_checkpoint_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#G1/S_checkpoint_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>G1/S checkpoint evolution</span> </div> </a> <ul id="toc-G1/S_checkpoint_evolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</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"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" 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">Cell cycle</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 58 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-58" 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">58 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%AF%D9%88%D8%B1%D8%A9_%D8%A7%D9%84%D8%AE%D9%84%D9%8A%D8%A9" title="دورة الخلية – Arabic" lang="ar" hreflang="ar" data-title="دورة الخلية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%9A%D2%AF%D2%99%D3%99%D0%BD%D3%99%D0%BA_%D1%86%D0%B8%D0%BA%D0%BB%D1%8B" title="Күҙәнәк циклы – Bashkir" lang="ba" hreflang="ba" data-title="Күҙәнәк циклы" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target"><span>Башҡортса</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%96%D0%B8%D0%B7%D0%BD%D0%B5%D0%BD_%D1%86%D0%B8%D0%BA%D1%8A%D0%BB_%D0%BD%D0%B0_%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="Жизнен цикъл на клетка – Bulgarian" lang="bg" hreflang="bg" data-title="Жизнен цикъл на клетка" 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/%C4%86elijski_ciklus" title="Ćelijski ciklus – Bosnian" lang="bs" hreflang="bs" data-title="Ćelijski ciklus" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/Kelc%27hiad_kelligel" title="Kelc'hiad kelligel – Breton" lang="br" hreflang="br" data-title="Kelc'hiad kelligel" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target"><span>Brezhoneg</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Cicle_cel%C2%B7lular" title="Cicle cel·lular – Catalan" lang="ca" hreflang="ca" data-title="Cicle cel·lular" 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/Bun%C4%9B%C4%8Dn%C3%BD_cyklus" title="Buněčný cyklus – Czech" lang="cs" hreflang="cs" data-title="Buněčný cyklus" 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/Zellzyklus" title="Zellzyklus – German" lang="de" hreflang="de" data-title="Zellzyklus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Rakuts%C3%BCkkel" title="Rakutsükkel – Estonian" lang="et" hreflang="et" data-title="Rakutsükkel" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CF%85%CF%84%CF%84%CE%B1%CF%81%CE%B9%CE%BA%CF%8C%CF%82_%CE%BA%CF%8D%CE%BA%CE%BB%CE%BF%CF%82" title="Κυτταρικός κύκλος – Greek" lang="el" hreflang="el" data-title="Κυτταρικός κύκλος" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Ciclo_celular" title="Ciclo celular – Spanish" lang="es" hreflang="es" data-title="Ciclo celular" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/%C4%88ela_ciklo" title="Ĉela ciklo – Esperanto" lang="eo" hreflang="eo" data-title="Ĉela ciklo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Zelula_ziklo" title="Zelula ziklo – Basque" lang="eu" hreflang="eu" data-title="Zelula ziklo" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%86%D8%B1%D8%AE%D9%87_%D8%B3%D9%84%D9%88%D9%84" title="چرخه سلول – Persian" lang="fa" hreflang="fa" data-title="چرخه سلول" 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/Cycle_cellulaire" title="Cycle cellulaire – French" lang="fr" hreflang="fr" data-title="Cycle cellulaire" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Ciclo_celular" title="Ciclo celular – Galician" lang="gl" hreflang="gl" data-title="Ciclo celular" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%84%B8%ED%8F%AC_%EC%A3%BC%EA%B8%B0" title="세포 주기 – Korean" lang="ko" hreflang="ko" data-title="세포 주기" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D5%BB%D5%BB%D5%AB_%D5%AF%D5%A5%D5%B6%D5%BD%D5%A1%D6%81%D5%AB%D5%AF%D5%AC" title="Բջջի կենսացիկլ – Armenian" lang="hy" hreflang="hy" data-title="Բջջի կենսացիկլ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%95%E0%A5%8B%E0%A4%B6%E0%A4%BF%E0%A4%95%E0%A4%BE_%E0%A4%9A%E0%A4%95%E0%A5%8D%E0%A4%B0" title="कोशिका चक्र – Hindi" lang="hi" hreflang="hi" data-title="कोशिका चक्र" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Stani%C4%8Dni_ciklus" title="Stanični ciklus – Croatian" lang="hr" hreflang="hr" data-title="Stanični ciklus" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Siklus_sel" title="Siklus sel – Indonesian" lang="id" hreflang="id" data-title="Siklus sel" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Frumuhringur" title="Frumuhringur – Icelandic" lang="is" hreflang="is" data-title="Frumuhringur" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Ciclo_cellulare" title="Ciclo cellulare – Italian" lang="it" hreflang="it" data-title="Ciclo cellulare" 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%9E%D7%97%D7%96%D7%95%D7%A8_%D7%94%D7%AA%D7%90" title="מחזור התא – Hebrew" lang="he" hreflang="he" data-title="מחזור התא" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Siklus_Sel" title="Siklus Sel – Javanese" lang="jv" hreflang="jv" data-title="Siklus Sel" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%96%D0%B0%D1%81%D1%83%D1%88%D0%B0%D0%BD%D1%8B%D2%A3_%D1%82%D1%96%D1%80%D1%88%D1%96%D0%BB%D1%96%D0%BA_%D1%86%D0%B8%D0%BA%D0%BB%D1%96" title="Жасушаның тіршілік циклі – Kazakh" lang="kk" hreflang="kk" data-title="Жасушаның тіршілік циклі" 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/Sik_selil%C3%A8" title="Sik selilè – Haitian Creole" lang="ht" hreflang="ht" data-title="Sik selilè" 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-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/%C3%87erxa_xaney%C3%AA" title="Çerxa xaneyê – Kurdish" lang="ku" hreflang="ku" data-title="Çerxa xaneyê" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Cyclus_cellularis" title="Cyclus cellularis – Latin" lang="la" hreflang="la" data-title="Cyclus cellularis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Sejtciklus" title="Sejtciklus – Hungarian" lang="hu" hreflang="hu" data-title="Sejtciklus" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Kitaran_sel" title="Kitaran sel – Malay" lang="ms" hreflang="ms" data-title="Kitaran sel" 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-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%AD%D1%81%D0%B8%D0%B9%D0%BD_%D0%BE%D1%80%D1%87%D0%B8%D0%BB" title="Эсийн орчил – Mongolian" lang="mn" hreflang="mn" data-title="Эсийн орчил" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Celcyclus" title="Celcyclus – Dutch" lang="nl" hreflang="nl" data-title="Celcyclus" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E5%91%A8%E6%9C%9F" title="細胞周期 – Japanese" lang="ja" hreflang="ja" data-title="細胞周期" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Cellesyklus" title="Cellesyklus – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Cellesyklus" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Cellesyklus" title="Cellesyklus – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Cellesyklus" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-om mw-list-item"><a href="https://om.wikipedia.org/wiki/Marsaa_seelii" title="Marsaa seelii – Oromo" lang="om" hreflang="om" data-title="Marsaa seelii" 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/Hujayra_sikli" title="Hujayra sikli – Uzbek" lang="uz" hreflang="uz" data-title="Hujayra sikli" 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/Cykl_kom%C3%B3rkowy" title="Cykl komórkowy – Polish" lang="pl" hreflang="pl" data-title="Cykl komórkowy" 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/Ciclo_celular" title="Ciclo celular – Portuguese" lang="pt" hreflang="pt" data-title="Ciclo celular" 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/Ciclu_celular" title="Ciclu celular – Romanian" lang="ro" hreflang="ro" data-title="Ciclu celular" 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%9A%D0%BB%D0%B5%D1%82%D0%BE%D1%87%D0%BD%D1%8B%D0%B9_%D1%86%D0%B8%D0%BA%D0%BB" title="Клеточный цикл – Russian" lang="ru" hreflang="ru" data-title="Клеточный цикл" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple badge-Q70893996 mw-list-item" title=""><a href="https://simple.wikipedia.org/wiki/Cell_cycle" title="Cell cycle – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cell cycle" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Bunkov%C3%BD_cyklus" title="Bunkový cyklus – Slovak" lang="sk" hreflang="sk" data-title="Bunkový cyklus" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Celi%C4%8Dni_ciklus" title="Celični ciklus – Slovenian" lang="sl" hreflang="sl" data-title="Celični ciklus" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%B3%D9%88%D9%88%DA%95%DB%8C_%D8%AE%D8%A7%D9%86%DB%95" title="سووڕی خانە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="سووڕی خانە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%96%D0%B8%D0%B2%D0%BE%D1%82%D0%BD%D0%B8_%D1%86%D0%B8%D0%BA%D0%BB%D1%83%D1%81_%D1%9B%D0%B5%D0%BB%D0%B8%D1%98%D0%B5" title="Животни циклус ћелије – Serbian" lang="sr" hreflang="sr" data-title="Животни циклус ћелије" 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/%C5%BDivotni_ciklus_%C4%87elije" title="Životni ciklus ćelije – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Životni ciklus ćelije" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Solusykli" title="Solusykli – Finnish" lang="fi" hreflang="fi" data-title="Solusykli" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Cellcykeln" title="Cellcykeln – Swedish" lang="sv" hreflang="sv" data-title="Cellcykeln" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Siklo_ng_selula" title="Siklo ng selula – Tagalog" lang="tl" hreflang="tl" data-title="Siklo ng selula" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%95%E0%AE%B2_%E0%AE%B5%E0%AE%9F%E0%AF%8D%E0%AE%9F%E0%AE%AE%E0%AF%8D" title="கல வட்டம் – Tamil" lang="ta" hreflang="ta" data-title="கல வட்டம்" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A7%E0%B8%B1%E0%B8%8F%E0%B8%88%E0%B8%B1%E0%B8%81%E0%B8%A3%E0%B9%80%E0%B8%8B%E0%B8%A5%E0%B8%A5%E0%B9%8C" title="วัฏจักรเซลล์ – Thai" lang="th" hreflang="th" data-title="วัฏจักรเซลล์" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/H%C3%BCcre_d%C3%B6ng%C3%BCs%C3%BC" title="Hücre döngüsü – Turkish" lang="tr" hreflang="tr" data-title="Hücre döngüsü" 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%9A%D0%BB%D1%96%D1%82%D0%B8%D0%BD%D0%BD%D0%B8%D0%B9_%D1%86%D0%B8%D0%BA%D0%BB" title="Клітинний цикл – Ukrainian" lang="uk" hreflang="uk" data-title="Клітинний цикл" 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/Chu_k%E1%BB%B3_t%E1%BA%BF_b%C3%A0o" title="Chu kỳ tế bào – Vietnamese" lang="vi" hreflang="vi" data-title="Chu kỳ tế bào" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E9%80%B1%E6%9C%9F" title="細胞週期 – Cantonese" lang="yue" hreflang="yue" data-title="細胞週期" data-language-autonym="粵語" data-language-local-name="Cantonese" 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class="noprint">From Wikipedia, the 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">Series of events and stages that result in cell division</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about the eukaryotic cell cycle. For the prokaryotic cell cycle, see <a href="/wiki/Fission_(biology)" title="Fission (biology)">fission (biology)</a>. For the separation of chromosomes that occurs as part of the cell cycle, see <a href="/wiki/Mitosis" title="Mitosis">mitosis</a>. For the academic journal, see <a href="/wiki/Cell_Cycle_(journal)" class="mw-redirect" title="Cell Cycle (journal)"><i>Cell Cycle</i> (journal)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Cell_division" title="Cell division">Cell division</a></div> <p class="mw-empty-elt"> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Normal_Cell_Life_Cycle.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Normal_Cell_Life_Cycle.png/200px-Normal_Cell_Life_Cycle.png" decoding="async" width="200" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Normal_Cell_Life_Cycle.png/300px-Normal_Cell_Life_Cycle.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Normal_Cell_Life_Cycle.png/400px-Normal_Cell_Life_Cycle.png 2x" data-file-width="800" data-file-height="600" /></a><figcaption>Life cycle of the cell</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Wilson1900Fig2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Wilson1900Fig2.jpg/250px-Wilson1900Fig2.jpg" decoding="async" width="200" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Wilson1900Fig2.jpg/330px-Wilson1900Fig2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/Wilson1900Fig2.jpg/500px-Wilson1900Fig2.jpg 2x" data-file-width="2211" data-file-height="1545" /></a><figcaption>Onion (<i><a href="/wiki/Allium" title="Allium">Allium</a></i>) cells in different phases of the cell cycle. Growth in an '<a href="/wiki/Organism" title="Organism">organism</a>' is carefully controlled by regulating the cell cycle.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/Reconstitution_in_6_phases%2C_of_the_changes_in_cell_shape_%28in_red%29_and_nucleoid_structure_%28roughly_equivalent_to_the_DNA%2C_in_green%29%2C_during_the_cell_cycle_of_the_bacteria_D._radiodurans.webm/220px--thumbnail.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="124" data-durationhint="61" data-mwtitle="Reconstitution_in_6_phases,_of_the_changes_in_cell_shape_(in_red)_and_nucleoid_structure_(roughly_equivalent_to_the_DNA,_in_green),_during_the_cell_cycle_of_the_bacteria_D._radiodurans.webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Reconstitution_in_6_phases,_of_the_changes_in_cell_shape_(in_red)_and_nucleoid_structure_(roughly_equivalent_to_the_DNA,_in_green),_during_the_cell_cycle_of_the_bacteria_D._radiodurans.webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/8/85/Reconstitution_in_6_phases%2C_of_the_changes_in_cell_shape_%28in_red%29_and_nucleoid_structure_%28roughly_equivalent_to_the_DNA%2C_in_green%29%2C_during_the_cell_cycle_of_the_bacteria_D._radiodurans.webm/Reconstitution_in_6_phases%2C_of_the_changes_in_cell_shape_%28in_red%29_and_nucleoid_structure_%28roughly_equivalent_to_the_DNA%2C_in_green%29%2C_during_the_cell_cycle_of_the_bacteria_D._radiodurans.webm.480p.vp9.webm" type="video/webm; 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codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /></video></span><figcaption>Cell cycle in <i>Deinococcus radiodurans</i></figcaption></figure> <p>The <b>cell cycle</b>, or <b>cell-division cycle</b>, is the sequential series of events that take place in a <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> that causes it to divide into two daughter cells. These events include the growth of the cell, duplication of its DNA (<a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>) and some of its <a href="/wiki/Organelle" title="Organelle">organelles</a>, and subsequently the partitioning of its cytoplasm, chromosomes and other components into two daughter cells in a process called <a href="/wiki/Cell_division" title="Cell division">cell division</a>. </p><p>In <a href="/wiki/Eukaryotic_cells" class="mw-redirect" title="Eukaryotic cells">eukaryotic cells</a> (having a <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>) including <a href="/wiki/Animal" title="Animal">animal</a>, <a href="/wiki/Plant" title="Plant">plant</a>, <a href="/wiki/Fungal" class="mw-redirect" title="Fungal">fungal</a>, and <a href="/wiki/Protist" title="Protist">protist</a> cells, the cell cycle is divided into two main stages: <a href="/wiki/Interphase" title="Interphase">interphase</a>, and the <a href="/wiki/M_phase" class="mw-redirect" title="M phase">M phase</a> that includes mitosis and cytokinesis.<sup id="cite_ref-Alberts2019_1-0" class="reference"><a href="#cite_note-Alberts2019-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> During interphase, the cell grows, accumulating nutrients needed for mitosis, and replicates its DNA and some of its organelles. During the M phase, the replicated <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>, organelles, and cytoplasm separate into two new daughter cells. To ensure the proper replication of cellular components and division, there are control mechanisms known as <a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">cell cycle checkpoints</a> after each of the key steps of the cycle that determine if the cell can progress to the next phase. </p><p>In cells without nuclei the <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>, <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>, the <a href="/wiki/Fission_(biology)" title="Fission (biology)">cell cycle</a> is divided into the B, C, and D periods. The B period extends from the end of cell division to the beginning of DNA replication. DNA replication occurs during the C period. The D period refers to the stage between the end of DNA replication and the splitting of the bacterial cell into two daughter cells.<sup id="cite_ref-Wang2009_2-0" class="reference"><a href="#cite_note-Wang2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>In single-celled organisms, a single cell-division cycle is how the organism reproduces to ensure its survival. In multicellular organisms such as plants and animals, a series of cell-division cycles is how the organism develops from a single-celled <a href="/wiki/Fertilized_egg" class="mw-redirect" title="Fertilized egg">fertilized egg</a> into a mature organism, and is also the process by which <a href="/wiki/Hair" title="Hair">hair</a>, <a href="/wiki/Skin" title="Skin">skin</a>, <a href="/wiki/Blood_cell" title="Blood cell">blood cells</a>, and some <a href="/wiki/Viscus" class="mw-redirect" title="Viscus">internal organs</a> are <a href="/wiki/Regeneration_(biology)" title="Regeneration (biology)">regenerated</a> and <a href="/wiki/Healing" title="Healing">healed</a> (with possible exception of <a href="/wiki/Nerve" title="Nerve">nerves</a>; see <a href="/wiki/Nerve_injury" title="Nerve injury">nerve damage</a>). After cell division, each of the daughter cells begin the <a href="/wiki/Interphase" title="Interphase">interphase</a> of a new cell cycle. Although the various stages of interphase are not usually morphologically distinguishable, each phase of the cell cycle has a distinct set of specialized biochemical processes that prepare the cell for initiation of the cell division. </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=Cell_cycle&action=edit&section=1" title="Edit section: Phases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The eukaryotic cell cycle consists of four distinct phases: <a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub> phase</a>, <a href="/wiki/S_phase" title="S phase">S phase</a> (synthesis), <a href="/wiki/G2_phase" title="G2 phase">G<sub>2</sub> phase</a> (collectively known as <a href="/wiki/Interphase" title="Interphase">interphase</a>) and <a href="/wiki/Mitosis" title="Mitosis">M phase</a> (mitosis and cytokinesis). M phase is itself composed of two tightly coupled processes: mitosis, in which the cell's nucleus divides, and <a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinesis</a>, in which the cell's <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> and cell membrane divides forming two daughter cells. Activation of each phase is dependent on the proper progression and completion of the previous one. Cells that have temporarily or reversibly stopped dividing are said to have entered a state of quiescence known as <a href="/wiki/G0_phase" title="G0 phase">G<sub>0</sub> phase</a> or <i>resting phase</i>. </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Cell_Cycle_2-2.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cell_Cycle_2-2.svg/300px-Cell_Cycle_2-2.svg.png" decoding="async" width="300" height="306" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cell_Cycle_2-2.svg/450px-Cell_Cycle_2-2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cell_Cycle_2-2.svg/600px-Cell_Cycle_2-2.svg.png 2x" data-file-width="748" data-file-height="764" /></a><figcaption>Schematic of the cell cycle. Outer ring: I = <a href="/wiki/Interphase" title="Interphase">Interphase</a>, M = <a href="/wiki/Mitosis" title="Mitosis">Mitosis</a>; inner ring: M = <a href="/wiki/Mitosis" title="Mitosis">Mitosis</a>, G<sub>1</sub> = <a href="/wiki/G1_phase" title="G1 phase">Gap 1</a>, G<sub>2</sub> = <a href="/wiki/G2_phase" title="G2 phase">Gap 2</a>, S = <a href="/wiki/S_phase" title="S phase">Synthesis</a>; not in ring: G<sub>0</sub> = <a href="/wiki/G0_phase" title="G0 phase">Gap 0/Resting</a><sup id="cite_ref-isbn0-87893-106-6_3-0" class="reference"><a href="#cite_note-isbn0-87893-106-6-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></figcaption></figure> <table class="wikitable" style="text-align:center"> <tbody><tr> <th>State </th> <th>Phase </th> <th>Abbreviation </th> <th>Description </th></tr> <tr> <td>Resting </td> <td style="height:50px"><a href="/wiki/G0_phase" title="G0 phase">Gap 0</a> </td> <td><b>G<sub>0</sub></b> </td> <td align="left" width="500pt">A phase where the cell has left the cycle and has stopped dividing. </td></tr> <tr> <td rowspan="3"><a href="/wiki/Interphase" title="Interphase">Interphase</a> </td> <td style="height:50px"><a href="/wiki/G1_phase" title="G1 phase">Gap 1</a> </td> <td><b>G<sub>1</sub></b> </td> <td align="left" width="500pt">Cell growth. The <i><a href="/wiki/Cell_cycle_checkpoint#G1_.28Restriction.29_Checkpoint" title="Cell cycle checkpoint">G<sub>1</sub> checkpoint</a></i> ensures that everything is ready for <a href="/wiki/DNA" title="DNA">DNA</a> synthesis. </td></tr> <tr> <td style="height:50px"><a href="/wiki/S_phase" title="S phase">Synthesis</a> </td> <td><b>S</b> </td> <td align="left" width="500pt"><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. </td></tr> <tr> <td style="height:50px"><a href="/wiki/G2_phase" title="G2 phase">Gap 2</a> </td> <td><b>G<sub>2</sub></b> </td> <td align="left" width="500pt">Growth and preparation for mitosis. The <i><a href="/wiki/Cell_cycle_checkpoint#G2_Checkpoint" title="Cell cycle checkpoint">G<sub>2</sub> checkpoint</a></i> ensures that everything is ready to enter the M (mitosis) phase and divide. </td></tr> <tr> <td><a href="/wiki/Cell_division" title="Cell division">Cell division</a> </td> <td style="height:50px"><a href="/wiki/Mitosis" title="Mitosis">Mitosis</a> </td> <td><b>M</b> </td> <td align="left" width="500pt">Cell division occurs. The <i><a href="/wiki/Cell_cycle_checkpoint#Metaphase_Checkpoint" title="Cell cycle checkpoint">Metaphase Checkpoint</a></i> ensures that the cell is ready to complete cell division. </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="G0_phase_(quiescence)"><span id="G0_phase_.28quiescence.29"></span>G<sub>0</sub> phase (quiescence)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=2" title="Edit section: G0 phase (quiescence)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Plant_cell_cycle.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Plant_cell_cycle.svg/220px-Plant_cell_cycle.svg.png" decoding="async" width="220" height="189" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Plant_cell_cycle.svg/330px-Plant_cell_cycle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Plant_cell_cycle.svg/440px-Plant_cell_cycle.svg.png 2x" data-file-width="1440" data-file-height="1240" /></a><figcaption>Plant cell cycle</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Animal_cell_cycle-en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Animal_cell_cycle-en.svg/220px-Animal_cell_cycle-en.svg.png" decoding="async" width="220" height="214" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Animal_cell_cycle-en.svg/330px-Animal_cell_cycle-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Animal_cell_cycle-en.svg/440px-Animal_cell_cycle-en.svg.png 2x" data-file-width="7207" data-file-height="7003" /></a><figcaption>Animal cell cycle</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/G0_phase" title="G0 phase">G0 phase</a></div> <p>G<sub>0</sub> is a resting phase where the cell has left the cycle and has stopped dividing. The cell cycle starts with this phase. Non-proliferative (non-dividing) cells in multicellular <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> generally enter the quiescent G<sub>0</sub> state from G<sub>1</sub> and may remain quiescent for long periods of time, possibly indefinitely (as is often the case for <a href="/wiki/Neuron" title="Neuron">neurons</a>). This is very common for cells that are fully <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiated</a>. Some cells enter the G<sub>0</sub> phase semi-permanently and are considered post-mitotic, e.g., some liver, kidney, and stomach cells. Many cells do not enter G<sub>0</sub> and continue to divide throughout an organism's life, e.g., epithelial cells. </p><p>The word "post-mitotic" is sometimes used to refer to both <a href="/wiki/G0_phase" title="G0 phase">quiescent</a> and <a href="/wiki/Cellular_senescence" title="Cellular senescence">senescent</a> cells. Cellular senescence occurs in response to DNA damage and external stress and usually constitutes an arrest in G<sub>1</sub>. Cellular senescence may make a cell's progeny nonviable; it is often a biochemical alternative to the self-destruction of such a damaged cell by <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Interphase">Interphase</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=3" title="Edit section: Interphase"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Interphase" title="Interphase">Interphase</a></div> <p>Interphase represents the phase between two successive M phases. Interphase is a series of changes that takes place in a newly formed cell and its nucleus before it becomes capable of division again. It is also called preparatory phase or intermitosis. Typically interphase lasts for at least 91% of the total time required for the cell cycle. </p><p>Interphase proceeds in three stages, G<sub>1</sub>, S, and G<sub>2</sub>, followed by the cycle of mitosis and cytokinesis. The cell's nuclear DNA contents are duplicated during S phase. </p> <div class="mw-heading mw-heading4"><h4 id="G1_phase_(First_growth_phase_or_Post_mitotic_gap_phase)"><span id="G1_phase_.28First_growth_phase_or_Post_mitotic_gap_phase.29"></span>G<sub>1</sub> phase (First growth phase or Post mitotic gap phase)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=4" title="Edit section: G1 phase (First growth phase or Post mitotic gap phase)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Human_karyotype_with_bands_and_sub-bands.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/250px-Human_karyotype_with_bands_and_sub-bands.png" decoding="async" width="250" height="413" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/375px-Human_karyotype_with_bands_and_sub-bands.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/500px-Human_karyotype_with_bands_and_sub-bands.png 2x" data-file-width="9684" data-file-height="16008" /></a><figcaption>Schematic <a href="/wiki/Karyogram" class="mw-redirect" title="Karyogram">karyogram</a> of the human chromosomes, showing their usual state in the G<sub>0</sub> and G<sub>1</sub> phase of the cell cycle. At top center it also shows the chromosome 3 pair in <a href="/wiki/Metaphase" title="Metaphase">metaphase</a> (annotated as "Meta."), which takes place after having undergone <a href="/wiki/DNA_synthesis" title="DNA synthesis">DNA synthesis</a> which occurs in the <a href="/wiki/S_phase" title="S phase">S phase</a> (annotated as S) of the cell cycle.<br /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Karyotype" title="Karyotype">Karyotype</a></div></figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a></div> <p>The first phase within interphase, from the end of the previous M phase until the beginning of DNA synthesis, is called <a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub></a> (G indicating <i>gap</i>). It is also called the growth phase. During this phase, the biosynthetic activities of the cell, which are considerably slowed down during M phase, resume at a high rate. The duration of G<sub>1</sub> is highly variable, even among different cells of the same species.<sup id="cite_ref-pmid4515625_4-0" class="reference"><a href="#cite_note-pmid4515625-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In this phase, the cell increases its supply of proteins, increases the number of organelles (such as mitochondria, ribosomes), and grows in size. In G<sub>1</sub> phase, a cell has three options. </p> <ul><li>To continue cell cycle and enter <a href="/wiki/S_phase" title="S phase">S phase</a></li> <li>Stop cell cycle and enter <a href="/wiki/G0_phase" title="G0 phase">G<sub>0</sub></a> phase for undergoing <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a>.</li> <li>Become arrested in G<sub>1</sub> phase hence it may enter G<sub>0</sub> phase or re-enter cell cycle.</li></ul> <p>The deciding point is called <a href="/wiki/Cell_cycle_checkpoint#G1_.28Restriction.29_Checkpoint" title="Cell cycle checkpoint">check point</a> (<a href="/wiki/Restriction_point" title="Restriction point">Restriction point</a>). This <a href="/wiki/Cell_cycle_checkpoint#G1_.28Restriction.29_Checkpoint" title="Cell cycle checkpoint">check point</a> is called the restriction point or START and is regulated by G<sub>1</sub>/S cyclins, which cause transition from G<sub>1</sub> to S phase. Passage through the G<sub>1</sub> check point commits the cell to division. </p> <div class="mw-heading mw-heading4"><h4 id="S_phase_(DNA_replication)"><span id="S_phase_.28DNA_replication.29"></span>S phase (DNA replication)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=5" title="Edit section: S phase (DNA replication)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/S_phase" title="S phase">S phase</a></div> <p>The ensuing <a href="/wiki/S_phase" title="S phase">S phase</a> starts when <a href="/wiki/DNA_synthesis" title="DNA synthesis">DNA synthesis</a> commences; when it is complete, all of the <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> have been replicated, i.e., each chromosome consists of two sister <a href="/wiki/Chromatid" title="Chromatid">chromatids</a>. Thus, during this phase, the amount of DNA in the cell has doubled, though the <a href="/wiki/Ploidy" title="Ploidy">ploidy</a> and number of chromosomes are unchanged. Rates of RNA <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> and <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein synthesis</a> are very low during this phase. An exception to this is <a href="/wiki/Histone" title="Histone">histone</a> production, most of which occurs during the S phase.<sup id="cite_ref-pmid7199388_5-0" class="reference"><a href="#cite_note-pmid7199388-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid12370293_6-0" class="reference"><a href="#cite_note-pmid12370293-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid14018040_7-0" class="reference"><a href="#cite_note-pmid14018040-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="G2_phase_(growth)"><span id="G2_phase_.28growth.29"></span>G<sub>2</sub> phase (growth)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=6" title="Edit section: G2 phase (growth)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/G2_phase" title="G2 phase">G2 phase</a></div> <p>G<sub>2</sub> phase occurs after DNA replication and is a period of protein synthesis and rapid cell growth to prepare the cell for mitosis. During this phase microtubules begin to reorganize to form a spindle (preprophase). Before proceeding to <a href="/wiki/Mitosis" title="Mitosis">mitotic phase</a>, cells must be checked at the G<sub>2</sub> checkpoint for any DNA damage within the chromosomes. The G<sub>2</sub> checkpoint is mainly regulated by the tumor protein <a href="/wiki/P53_upregulated_modulator_of_apoptosis" title="P53 upregulated modulator of apoptosis">p53</a>. If the DNA is damaged, p53 will either repair the DNA or trigger the apoptosis of the cell. If p53 is dysfunctional or mutated, cells with damaged DNA may continue through the cell cycle, leading to the development of cancer. </p> <div class="mw-heading mw-heading3"><h3 id="Mitotic_phase_(chromosome_separation)"><span id="Mitotic_phase_.28chromosome_separation.29"></span>Mitotic phase (chromosome separation)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=7" title="Edit section: Mitotic phase (chromosome separation)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Mitosis" title="Mitosis">Mitosis</a></div> <p>The relatively brief <i>M phase</i> consists of nuclear division (<a href="/wiki/Karyokinesis" class="mw-redirect" title="Karyokinesis">karyokinesis</a>) and division of cytoplasm (<a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinesis</a>). M phase is complex and highly regulated. The sequence of events is divided into phases, corresponding to the completion of one set of activities and the start of the next. These phases are sequentially known as: </p> <ul><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/Anaphase" title="Anaphase">anaphase</a></li> <li><a href="/wiki/Telophase" title="Telophase">telophase</a></li></ul> <figure class="mw-halign-center" typeof="mw:File"><a href="/wiki/File:Mitosis_Stages.svg" class="mw-file-description" title="A diagram of the mitotic phases"><img alt="A diagram of the mitotic phases" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Mitosis_Stages.svg/900px-Mitosis_Stages.svg.png" decoding="async" width="900" height="156" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Mitosis_Stages.svg/1350px-Mitosis_Stages.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c9/Mitosis_Stages.svg/1800px-Mitosis_Stages.svg.png 2x" data-file-width="2361" data-file-height="409" /></a><figcaption>A diagram of the mitotic phases</figcaption></figure> <p>Mitosis is the process by which a <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> cell separates the <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> in its <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a> into two identical sets in two nuclei.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> During the process of mitosis the pairs of <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a> condense and attach to <a href="/wiki/Microtubule" title="Microtubule">microtubules</a> that pull the <a href="/wiki/Sister_chromatids" title="Sister chromatids">sister chromatids</a> to opposite sides of the cell.<sup id="cite_ref-Maton_1997_9-0" class="reference"><a href="#cite_note-Maton_1997-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>Mitosis occurs exclusively in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells, but occurs in different ways in different species. For example, animal cells undergo an "open" mitosis, where the <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a> breaks down before the chromosomes separate, while <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> such as <i><a href="/wiki/Aspergillus_nidulans" title="Aspergillus nidulans">Aspergillus nidulans</a></i> and <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> (<a href="/wiki/Yeast" title="Yeast">yeast</a>) undergo a "closed" mitosis, where chromosomes divide within an intact <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cytokinesis_phase_(separation_of_all_cell_components)"><span id="Cytokinesis_phase_.28separation_of_all_cell_components.29"></span>Cytokinesis phase (separation of all cell components)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=8" title="Edit section: Cytokinesis phase (separation of all cell components)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cytokinesis" title="Cytokinesis">Cytokinesis</a></div> <p>Mitosis is immediately followed by <a href="/wiki/Cytokinesis" title="Cytokinesis">cytokinesis</a>, which divides the nuclei, <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>, <a href="/wiki/Organelle" title="Organelle">organelles</a> and <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> into two cells containing roughly equal shares of these cellular components. Cytokinesis occurs differently in plant and animal cells. While the cell membrane forms a groove that gradually deepens to separate the cytoplasm in animal cells, a <a href="/wiki/Cell_plate" title="Cell plate">cell plate</a> is formed to separate it in plant cells. The position of the cell plate is determined by the position of a preprophase band of microtubules and <a href="/wiki/Actin" title="Actin">actin</a> filaments. Mitosis and cytokinesis together define the <a href="/wiki/Cell_division" title="Cell division">division</a> of the parent cell into two daughter cells, genetically identical to each other and to their parent cell. This accounts for approximately 10% of the cell cycle. </p><p>Because cytokinesis usually occurs in conjunction with mitosis, "mitosis" is often used interchangeably with "M phase". However, there are many cells where mitosis and cytokinesis occur separately, forming single cells with multiple nuclei in a process called <a href="/wiki/Endoreplication" class="mw-redirect" title="Endoreplication">endoreplication</a>. This occurs most notably among the <a href="/wiki/Fungus" title="Fungus">fungi</a> and <a href="/wiki/Slime_mold" title="Slime mold">slime molds</a>, but is found in various groups. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">fruit fly</a> embryonic development.<sup id="cite_ref-Lilly_11-0" class="reference"><a href="#cite_note-Lilly-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Errors in mitosis can result in cell death through <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> or cause <a href="/wiki/Mutation" title="Mutation">mutations</a> that may lead to <a href="/wiki/Cancer" title="Cancer">cancer</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Regulation_of_eukaryotic_cell_cycle"><span class="anchor" id="Regulation_of_cell_cycle"></span>Regulation of eukaryotic cell cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=9" title="Edit section: Regulation of eukaryotic cell cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:A_simplified_view_of_the_cell-cycle_control_system.pdf" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/A_simplified_view_of_the_cell-cycle_control_system.pdf/page1-220px-A_simplified_view_of_the_cell-cycle_control_system.pdf.jpg" decoding="async" width="220" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/A_simplified_view_of_the_cell-cycle_control_system.pdf/page1-330px-A_simplified_view_of_the_cell-cycle_control_system.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5d/A_simplified_view_of_the_cell-cycle_control_system.pdf/page1-440px-A_simplified_view_of_the_cell-cycle_control_system.pdf.jpg 2x" data-file-width="1125" data-file-height="487" /></a><figcaption>Levels of the three major cyclin types oscillate during the cell cycle (top), providing the basis for oscillations in the cyclin–Cdk complexes that drive cell-cycle events (bottom). In general, Cdk levels are constant and in large excess over cyclin levels; thus, cyclin–Cdk complexes form in parallel with cyclin levels. The enzymatic activities of cyclin–Cdk complexes also tend to rise and fall in parallel with cyclin levels, although in some cases Cdk inhibitor proteins or phosphorylation introduce a delay between the formation and activation of cyclin–Cdk complexes. Formation of active G1/S–Cdk complexes commits the cell to a new division cycle at the Start checkpoint in late G1. G1/S–Cdks then activate the S–Cdk complexes that initiate DNA replication at the beginning of S phase. M–Cdk activation occurs after the completion of S phase, resulting in progression through the G2/M checkpoint and assembly of the mitotic spindle. APC activation then triggers sister-chromatid separation at the metaphase-to-anaphase transition. APC activity also causes the destruction of S and M cyclins and thus the inactivation of Cdks, which promotes the completion of mitosis and cytokinesis. APC activity is maintained in G1 until G1/S–Cdk activity rises again and commits the cell to the next cycle. This scheme serves only as a general guide and does not apply to all cell types.</figcaption></figure> <p>Regulation of the cell cycle involves processes crucial to the survival of a cell, including the detection and repair of genetic damage as well as the prevention of uncontrolled cell division. The molecular events that control the cell cycle are ordered and directional; that is, each process occurs in a sequential fashion and it is impossible to "reverse" the cycle. </p> <div class="mw-heading mw-heading3"><h3 id="Role_of_cyclins_and_CDKs">Role of cyclins and CDKs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=10" title="Edit section: Role of cyclins and CDKs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table class="wikitable" align="right"> <tbody><tr align="center"> <td><span typeof="mw:File"><a href="/wiki/File:Paul_Nurse_portrait.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Paul_Nurse_portrait.jpg/120px-Paul_Nurse_portrait.jpg" decoding="async" width="120" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Paul_Nurse_portrait.jpg/250px-Paul_Nurse_portrait.jpg 1.5x" data-file-width="1648" data-file-height="2172" /></a></span><br />Nobel Laureate<br /><a href="/wiki/Paul_Nurse" title="Paul Nurse">Paul Nurse</a> </td> <td><span typeof="mw:File"><a href="/wiki/File:Tim_hunt.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Tim_hunt.jpg/132px-Tim_hunt.jpg" decoding="async" width="132" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Tim_hunt.jpg/198px-Tim_hunt.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Tim_hunt.jpg/264px-Tim_hunt.jpg 2x" data-file-width="546" data-file-height="654" /></a></span><br />Nobel Laureate<br /><a href="/wiki/Tim_Hunt" title="Tim Hunt">Tim Hunt</a> </td></tr></tbody></table> <p>Two key classes of regulatory molecules, <a href="/wiki/Cyclin" title="Cyclin">cyclins</a> and <a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">cyclin-dependent kinases</a> (CDKs), determine a cell's progress through the cell cycle.<sup id="cite_ref-pmid7575488_12-0" class="reference"><a href="#cite_note-pmid7575488-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Leland_H._Hartwell" title="Leland H. Hartwell">Leland H. Hartwell</a>, <a href="/wiki/R._Timothy_Hunt" class="mw-redirect" title="R. Timothy Hunt">R. Timothy Hunt</a>, and <a href="/wiki/Paul_M._Nurse" class="mw-redirect" title="Paul M. Nurse">Paul M. Nurse</a> won the 2001 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> for their discovery of these central molecules.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Many of the genes encoding cyclins and CDKs are <a href="/wiki/Conservation_(genetics)" class="mw-redirect" title="Conservation (genetics)">conserved</a> among all eukaryotes, but in general, more complex organisms have more elaborate cell cycle control systems that incorporate more individual components. Many of the relevant genes were first identified by studying yeast, especially <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>;<sup id="cite_ref-pmid9843569_14-0" class="reference"><a href="#cite_note-pmid9843569-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> genetic nomenclature in yeast dubs many of these genes <i>cdc</i> (for "cell division cycle") followed by an identifying number, e.g. <i><a href="/wiki/Cdc25" title="Cdc25">cdc25</a></i> or <i><a href="/wiki/Cdc20" class="mw-redirect" title="Cdc20">cdc20</a></i>. </p><p>Cyclins form the regulatory subunits and CDKs the catalytic subunits of an activated <a href="/wiki/Heterodimer" class="mw-redirect" title="Heterodimer">heterodimer</a>; cyclins have no catalytic activity and CDKs are inactive in the absence of a partner cyclin. When activated by a bound cyclin, CDKs perform a common biochemical reaction called <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a> that activates or inactivates target proteins to orchestrate coordinated entry into the next phase of the cell cycle. Different cyclin-CDK combinations determine the downstream proteins targeted. CDKs are constitutively expressed in cells whereas cyclins are synthesised at specific stages of the cell cycle, in response to various molecular signals.<sup id="cite_ref-Robbins_15-0" class="reference"><a href="#cite_note-Robbins-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="General_mechanism_of_cyclin-CDK_interaction">General mechanism of cyclin-CDK interaction</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=11" title="Edit section: General mechanism of cyclin-CDK interaction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Upon receiving a pro-mitotic extracellular signal, G<sub>1</sub> <a href="/wiki/Cyclin-CDK" class="mw-redirect" title="Cyclin-CDK">cyclin-CDK</a> complexes become active to prepare the cell for S phase, promoting the expression of <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> that in turn promote the expression of S cyclins and of enzymes required for <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. The G<sub>1</sub> cyclin-CDK complexes also promote the degradation of molecules that function as S phase inhibitors by targeting them for <a href="/wiki/Ubiquitination" class="mw-redirect" title="Ubiquitination">ubiquitination</a>. Once a protein has been ubiquitinated, it is targeted for proteolytic degradation by the <a href="/wiki/Proteasome" title="Proteasome">proteasome</a>. Results from a study of E2F transcriptional dynamics at the single-cell level argue that the role of G1 cyclin-CDK activities, in particular cyclin D-CDK4/6, is to tune the timing rather than the commitment of cell cycle entry.<sup id="cite_ref-Dong,_P._2014_16-0" class="reference"><a href="#cite_note-Dong,_P._2014-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Active S cyclin-CDK complexes phosphorylate proteins that make up the <a href="/wiki/Pre-replication_complex" title="Pre-replication complex">pre-replication complexes</a> assembled during G<sub>1</sub> phase on DNA <a href="/wiki/Origin_of_replication" title="Origin of replication">replication origins</a>. The phosphorylation serves two purposes: to activate each already-assembled pre-replication complex, and to prevent new complexes from forming. This ensures that every portion of the cell's <a href="/wiki/Genome" title="Genome">genome</a> will be replicated once and only once. The reason for prevention of gaps in replication is fairly clear, because daughter cells that are missing all or part of crucial genes will die. However, for reasons related to <a href="/wiki/Gene_copy_number" class="mw-redirect" title="Gene copy number">gene copy number</a> effects, possession of extra copies of certain genes is also deleterious to the daughter cells. </p><p>Mitotic cyclin-CDK complexes, which are synthesized but inactivated during S and G<sub>2</sub> phases, promote the initiation of <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> by stimulating downstream proteins involved in chromosome condensation and <a href="/wiki/Mitotic_spindle" class="mw-redirect" title="Mitotic spindle">mitotic spindle</a> assembly. A critical complex activated during this process is a <a href="/wiki/Ubiquitin_ligase" title="Ubiquitin ligase">ubiquitin ligase</a> known as the <a href="/wiki/Anaphase-promoting_complex" title="Anaphase-promoting complex">anaphase-promoting complex</a> (APC), which promotes degradation of structural proteins associated with the chromosomal <a href="/wiki/Kinetochore" title="Kinetochore">kinetochore</a>. APC also targets the mitotic cyclins for degradation, ensuring that telophase and cytokinesis can proceed.<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-heading4"><h4 id="Specific_action_of_cyclin-CDK_complexes">Specific action of cyclin-CDK complexes</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=12" title="Edit section: Specific action of cyclin-CDK complexes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Cyclin_D" title="Cyclin D">Cyclin D</a> is the first cyclin produced in the cells that enter the cell cycle, in response to extracellular signals (e.g. <a href="/wiki/Growth_factor" title="Growth factor">growth factors</a>). Cyclin D levels stay low in resting cells that are not proliferating. Additionally, <a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4/6</a> and <a href="/wiki/Cyclin-dependent_kinase_2" title="Cyclin-dependent kinase 2">CDK2</a> are also inactive because CDK4/6 are bound by <a href="/wiki/INK4" title="INK4">INK4</a> family members (e.g., p16), limiting kinase activity. Meanwhile, CDK2 complexes are inhibited by the CIP/KIP proteins such as p21 and p27,<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> When it is time for a cell to enter the cell cycle, which is triggered by a mitogenic stimuli, levels of cyclin D increase. In response to this trigger, cyclin D binds to existing <a href="/wiki/Cyclin-dependent_kinase_4" title="Cyclin-dependent kinase 4">CDK4</a>/6, forming the active cyclin D-CDK4/6 complex. Cyclin D-CDK4/6 complexes in turn mono-phosphorylates the <a href="/wiki/Retinoblastoma" title="Retinoblastoma">retinoblastoma</a> susceptibility protein (<a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">Rb</a>) to pRb. The un-phosphorylated Rb tumour suppressor functions in inducing cell cycle exit and maintaining G0 arrest (senescence).<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>In the last few decades, a model has been widely accepted whereby pRB proteins are inactivated by cyclin D-Cdk4/6-mediated phosphorylation. Rb has 14+ potential phosphorylation sites. Cyclin D-Cdk 4/6 progressively phosphorylates Rb to hyperphosphorylated state, which triggers dissociation of pRB–<a href="/wiki/E2F" title="E2F">E2F</a> complexes, thereby inducing G1/S cell cycle gene expression and progression into S phase.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Scientific observations from a study have shown that Rb is present in three types of isoforms: (1) un-phosphorylated Rb in G0 state; (2) mono-phosphorylated Rb, also referred to as "hypo-phosphorylated' or 'partially' phosphorylated Rb in early G1 state; and (3) inactive hyper-phosphorylated Rb in late G1 state.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_23-0" class="reference"><a href="#cite_note-:0-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> In early G1 cells, mono-phosphorylated Rb exists as 14 different isoforms, one of each has distinct <a href="/wiki/E2F" title="E2F">E2F</a> binding affinity.<sup id="cite_ref-:0_23-1" class="reference"><a href="#cite_note-:0-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Rb has been found to associate with hundreds of different proteins<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> and the idea that different mono-phosphorylated Rb isoforms have different protein partners was very appealing.<sup id="cite_ref-pmid27401552_25-0" class="reference"><a href="#cite_note-pmid27401552-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> A later report confirmed that mono-phosphorylation controls Rb's association with other proteins and generates functional distinct forms of Rb.<sup id="cite_ref-Sanidas_26-0" class="reference"><a href="#cite_note-Sanidas-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> All different mono-phosphorylated Rb isoforms inhibit E2F transcriptional program and are able to arrest cells in G1-phase. Different mono-phosphorylated forms of Rb have distinct transcriptional outputs that are extended beyond E2F regulation.<sup id="cite_ref-Sanidas_26-1" class="reference"><a href="#cite_note-Sanidas-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>In general, the binding of pRb to E2F inhibits the E2F target gene expression of certain G1/S and S transition genes including <a href="/wiki/Cyclin_E" title="Cyclin E">E-type cyclins</a>. The partial phosphorylation of Rb de-represses the Rb-mediated suppression of E2F target gene expression, begins the expression of cyclin E. The molecular mechanism that causes the cell switched to cyclin E activation is currently not known, but as cyclin E levels rise, the active cyclin E-CDK2 complex is formed, bringing Rb to be inactivated by hyper-phosphorylation.<sup id="cite_ref-:0_23-2" class="reference"><a href="#cite_note-:0-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Hyperphosphorylated Rb is completely dissociated from E2F, enabling further expression of a wide range of E2F target genes are required for driving cells to proceed into S phase [1]. It has been identified that cyclin D-Cdk4/6 binds to a C-terminal alpha-helix region of Rb that is only distinguishable to cyclin D rather than other cyclins, <a href="/wiki/Cyclin_E" title="Cyclin E">cyclin E</a>, <a href="/wiki/Cyclin_A" title="Cyclin A">A</a> and <a href="/wiki/Cyclin_B" title="Cyclin B">B</a>.<sup id="cite_ref-:1_27-0" class="reference"><a href="#cite_note-:1-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> This observation based on the structural analysis of Rb phosphorylation supports that Rb is phosphorylated in a different level through multiple Cyclin-Cdk complexes. This also makes feasible the current model of a simultaneous switch-like inactivation of all mono-phosphorylated Rb isoforms through one type of Rb hyper-phosphorylation mechanism. In addition, mutational analysis of the cyclin D- Cdk 4/6 specific Rb C-terminal helix shows that disruptions of cyclin D-Cdk 4/6 binding to Rb prevents Rb phosphorylation, arrests cells in G1, and bolsters Rb's functions in tumor suppressor.<sup id="cite_ref-:1_27-1" class="reference"><a href="#cite_note-:1-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> This cyclin-Cdk driven cell cycle transitional mechanism governs a cell committed to the cell cycle that allows cell proliferation. A cancerous cell growth often accompanies with deregulation of Cyclin D-Cdk 4/6 activity. </p><p>The hyperphosphorylated Rb dissociates from the E2F/DP1/Rb complex (which was bound to the <a href="/wiki/E2F" title="E2F">E2F</a> responsive genes, effectively "blocking" them from transcription), activating E2F. Activation of E2F results in transcription of various genes like <a href="/wiki/Cyclin_E" title="Cyclin E">cyclin E</a>, <a href="/wiki/Cyclin_A" title="Cyclin A">cyclin A</a>, <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a>, <a href="/wiki/Thymidine_kinase" title="Thymidine kinase">thymidine kinase</a>, etc. Cyclin E thus produced binds to <a href="/wiki/Cyclin-dependent_kinase_2" title="Cyclin-dependent kinase 2">CDK2</a>, forming the cyclin E-CDK2 complex, which pushes the cell from G<sub>1</sub> to S phase (G<sub>1</sub>/S, which initiates the G<sub>2</sub>/M transition).<sup id="cite_ref-isbn0-12-324719-5_28-0" class="reference"><a href="#cite_note-isbn0-12-324719-5-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Cyclin_B" title="Cyclin B">Cyclin B</a>-cdk1 complex activation causes breakdown of <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a> and initiation of <a href="/wiki/Prophase" title="Prophase">prophase</a>, and subsequently, its deactivation causes the cell to exit mitosis.<sup id="cite_ref-Robbins_15-1" class="reference"><a href="#cite_note-Robbins-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> A quantitative study of E2F transcriptional dynamics at the single-cell level by using engineered fluorescent reporter cells provided a quantitative framework for understanding the control logic of cell cycle entry, challenging the canonical textbook model. Genes that regulate the amplitude of E2F accumulation, such as Myc, determine the commitment in cell cycle and S phase entry. G1 cyclin-CDK activities are not the driver of cell cycle entry. Instead, they primarily tune the timing of E2F increase, thereby modulating the pace of cell cycle progression.<sup id="cite_ref-Dong,_P._2014_16-1" class="reference"><a href="#cite_note-Dong,_P._2014-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inhibitors">Inhibitors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=13" title="Edit section: Inhibitors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Endogenous">Endogenous</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=14" title="Edit section: Endogenous"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Signal_transduction_pathways.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/300px-Signal_transduction_pathways.svg.png" decoding="async" width="300" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/450px-Signal_transduction_pathways.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Signal_transduction_pathways.svg/600px-Signal_transduction_pathways.svg.png 2x" data-file-width="1858" data-file-height="1364" /></a><figcaption>Overview of signal transduction pathways involved in <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, also known as "programmed cell death"</figcaption></figure> <p>Two families of genes, the <i>cip/kip</i> (<i>CDK interacting protein/Kinase inhibitory protein</i>) family and the INK4a/ARF (<i>In</i>hibitor of <i>K</i>inase 4/<i>A</i>lternative <i>R</i>eading <i>F</i>rame) family, prevent the progression of the cell cycle. Because these genes are instrumental in prevention of <a href="/wiki/Tumor" class="mw-redirect" title="Tumor">tumor</a> formation, they are known as <a href="/wiki/Tumor_suppressor" class="mw-redirect" title="Tumor suppressor">tumor suppressors</a>. </p><p>The <b><i>cip/kip</i> family</b> includes the genes <a href="/wiki/P21" title="P21">p21</a>, <a href="/wiki/P27_(gene)" class="mw-redirect" title="P27 (gene)">p27</a> and <a href="/wiki/P57_(gene)" class="mw-redirect" title="P57 (gene)">p57</a>. They halt the cell cycle in G<sub>1</sub> phase by binding to and inactivating cyclin-CDK complexes. p21 is activated by <a href="/wiki/P53" title="P53">p53</a> (which, in turn, is triggered by DNA damage e.g. due to radiation). p27 is activated by Transforming Growth Factor β (<a href="/wiki/TGF_%CE%B2" class="mw-redirect" title="TGF β">TGF β</a>), a growth inhibitor. </p><p>The <b>INK4a/ARF family</b> includes <a href="/wiki/P16_(gene)" class="mw-redirect" title="P16 (gene)">p16<sup>INK4a</sup></a>, which binds to CDK4 and arrests the cell cycle in G<sub>1</sub> phase, and <a href="/wiki/P14arf" title="P14arf">p14<sup>ARF</sup></a> which prevents p53 degradation. </p> <div class="mw-heading mw-heading4"><h4 id="Synthetic">Synthetic</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=15" title="Edit section: Synthetic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Synthetic inhibitors of <a href="/wiki/Cdc25" title="Cdc25">Cdc25</a> could also be useful for the arrest of cell cycle and therefore be useful as antineoplastic and anticancer agents.<sup id="cite_ref-ref_1_29-0" class="reference"><a href="#cite_note-ref_1-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p>Many human cancers possess the hyper-activated Cdk 4/6 activities.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Given the observations of cyclin D-Cdk 4/6 functions, inhibition of Cdk 4/6 should result in preventing a malignant tumor from proliferating. Consequently, scientists have tried to invent the synthetic Cdk4/6 inhibitor as Cdk4/6 has been characterized to be a therapeutic target for anti-tumor effectiveness. Three Cdk4/6 inhibitors – <a href="/wiki/Palbociclib" title="Palbociclib">palbociclib</a>, <a href="/wiki/Ribociclib" title="Ribociclib">ribociclib</a>, and <a href="/wiki/Abemaciclib" title="Abemaciclib">abemaciclib</a> – currently received FDA approval for clinical use to treat advanced-stage or <a href="/wiki/Metastatic_breast_cancer" title="Metastatic breast cancer">metastatic</a>, <a href="/wiki/Hormone_receptor_positive_breast_tumor" title="Hormone receptor positive breast tumor">hormone-receptor-positive</a> (HR-positive, HR+), <a href="/wiki/HER2_negative_breast_cancer" class="mw-redirect" title="HER2 negative breast cancer">HER2-negative</a> (HER2-) breast cancer.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bilgin_2017_32-0" class="reference"><a href="#cite_note-Bilgin_2017-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> For example, palbociclib is an orally active CDK4/6 inhibitor which has demonstrated improved outcomes for ER-positive/HER2-negative advanced breast cancer. The main side effect is <a href="/wiki/Neutropenia" title="Neutropenia">neutropenia</a> which can be managed by dose reduction.<sup id="cite_ref-Schmidt_2018_33-0" class="reference"><a href="#cite_note-Schmidt_2018-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p><p>Cdk4/6 targeted therapy will only treat cancer types where Rb is expressed. Cancer cells with loss of Rb have primary resistance to Cdk4/6 inhibitors. </p> <div class="mw-heading mw-heading3"><h3 id="Transcriptional_regulatory_network">Transcriptional regulatory network</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=16" title="Edit section: Transcriptional regulatory network"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Current evidence suggests that a semi-autonomous transcriptional network acts in concert with the CDK-cyclin machinery to regulate the cell cycle. Several gene expression studies in <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> have identified 800–1200 genes that change expression over the course of the cell cycle.<sup id="cite_ref-pmid9843569_14-1" class="reference"><a href="#cite_note-pmid9843569-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pramilaetal2006_34-0" class="reference"><a href="#cite_note-pramilaetal2006-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-orlandoeta1nature2008_35-0" class="reference"><a href="#cite_note-orlandoeta1nature2008-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> They are transcribed at high levels at specific points in the cell cycle, and remain at lower levels throughout the rest of the cycle. While the set of identified genes differs between studies due to the computational methods and criteria used to identify them, each study indicates that a large portion of yeast genes are temporally regulated.<sup id="cite_ref-deLichtenberg2005_36-0" class="reference"><a href="#cite_note-deLichtenberg2005-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p><p>Many periodically expressed genes are driven by <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> that are also periodically expressed. One screen of single-gene knockouts identified 48 transcription factors (about 20% of all non-essential transcription factors) that show cell cycle progression defects.<sup id="cite_ref-whiteetal2009_37-0" class="reference"><a href="#cite_note-whiteetal2009-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Genome-wide studies using high throughput technologies have identified the transcription factors that bind to the promoters of yeast genes, and correlating these findings with temporal expression patterns have allowed the identification of transcription factors that drive phase-specific gene expression.<sup id="cite_ref-pramilaetal2006_34-1" class="reference"><a href="#cite_note-pramilaetal2006-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-leeetal2002_38-0" class="reference"><a href="#cite_note-leeetal2002-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The expression profiles of these transcription factors are driven by the transcription factors that peak in the prior phase, and computational models have shown that a CDK-autonomous network of these transcription factors is sufficient to produce steady-state oscillations in gene expression).<sup id="cite_ref-orlandoeta1nature2008_35-1" class="reference"><a href="#cite_note-orlandoeta1nature2008-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-simonetal2001_39-0" class="reference"><a href="#cite_note-simonetal2001-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>Experimental evidence also suggests that gene expression can oscillate with the period seen in dividing wild-type cells independently of the CDK machinery. Orlando <i>et al.</i> used <a href="/wiki/Microarray" title="Microarray">microarrays</a> to measure the expression of a set of 1,271 genes that they identified as periodic in both wild type cells and cells lacking all S-phase and mitotic cyclins (<i>clb1,2,3,4,5,6</i>). Of the 1,271 genes assayed, 882 continued to be expressed in the cyclin-deficient cells at the same time as in the wild type cells, despite the fact that the cyclin-deficient cells arrest at the border between <a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub></a> and <a href="/wiki/S_phase" title="S phase">S phase</a>. However, 833 of the genes assayed changed behavior between the wild type and mutant cells, indicating that these genes are likely directly or indirectly regulated by the CDK-cyclin machinery. Some genes that continued to be expressed on time in the mutant cells were also expressed at different levels in the mutant and wild type cells. These findings suggest that while the transcriptional network may oscillate independently of the CDK-cyclin oscillator, they are coupled in a manner that requires both to ensure the proper timing of cell cycle events.<sup id="cite_ref-orlandoeta1nature2008_35-2" class="reference"><a href="#cite_note-orlandoeta1nature2008-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Other work indicates that <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a>, a post-translational modification, of cell cycle transcription factors by <a href="/wiki/Cdk1" class="mw-redirect" title="Cdk1">Cdk1</a> may alter the localization or activity of the transcription factors in order to tightly control timing of target genes.<sup id="cite_ref-whiteetal2009_37-1" class="reference"><a href="#cite_note-whiteetal2009-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-sidorova1995_40-0" class="reference"><a href="#cite_note-sidorova1995-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ubersaxetal2003_41-0" class="reference"><a href="#cite_note-ubersaxetal2003-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p><p>While oscillatory transcription plays a key role in the progression of the yeast cell cycle, the CDK-cyclin machinery operates independently in the early embryonic cell cycle. Before the <a href="/wiki/Midblastula_transition" class="mw-redirect" title="Midblastula transition">midblastula transition</a>, <a href="/wiki/Zygote" title="Zygote">zygotic</a> transcription does not occur and all needed proteins, such as the B-type cyclins, are translated from maternally loaded <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a>.<sup id="cite_ref-davidmorganbook2007_42-0" class="reference"><a href="#cite_note-davidmorganbook2007-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="DNA_replication_and_DNA_replication_origin_activity">DNA replication and DNA replication origin activity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=17" title="Edit section: DNA replication and DNA replication origin activity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Analyses of synchronized cultures of <i>Saccharomyces cerevisiae</i> under conditions that prevent DNA replication initiation without delaying cell cycle progression showed that origin licensing decreases the expression of genes with origins near their 3' ends, revealing that downstream origins can regulate the expression of upstream genes.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> This confirms previous predictions from mathematical modeling of a global causal coordination between DNA replication origin activity and mRNA expression,<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> and shows that mathematical modeling of DNA microarray data can be used to correctly predict previously unknown biological modes of regulation. </p> <div class="mw-heading mw-heading2"><h2 id="Checkpoints">Checkpoints</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=18" title="Edit section: Checkpoints"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">Cell cycle checkpoint</a></div> <p><a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">Cell cycle checkpoints</a> are used by the cell to monitor and regulate the progress of the cell cycle.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Checkpoints prevent cell cycle progression at specific points, allowing verification of necessary phase processes and repair of <a href="/wiki/DNA_damage" class="mw-redirect" title="DNA damage">DNA damage</a>. The cell cannot proceed to the next phase until checkpoint requirements have been met. Checkpoints typically consist of a network of regulatory proteins that monitor and dictate the progression of the cell through the different stages of the cell cycle. </p><p>It is estimated that in normal human cells about 1% of <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">single-strand DNA damages</a> are converted to about 50 endogenous DNA double-strand breaks per cell per cell cycle.<sup id="cite_ref-Vilenchik2003_48-0" class="reference"><a href="#cite_note-Vilenchik2003-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Although such double-strand breaks are usually <a href="/wiki/DNA_repair" title="DNA repair">repaired</a> with high fidelity, errors in their repair are considered to contribute significantly to the rate of cancer in humans.<sup id="cite_ref-Vilenchik2003_48-1" class="reference"><a href="#cite_note-Vilenchik2003-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p><p>There are several checkpoints to ensure that damaged or incomplete DNA is not passed on to daughter cells. Three main checkpoints exist: the G<sub>1</sub>/S checkpoint, the G<sub>2</sub>/M checkpoint and the metaphase (mitotic) checkpoint. Another checkpoint is the Go checkpoint, in which the cells are checked for maturity. If the cells fail to pass this checkpoint by not being ready yet, they will be discarded from dividing. </p><p>G<sub>1</sub>/S transition is a rate-limiting step in the cell cycle and is also known as <a href="/wiki/Restriction_point" title="Restriction point">restriction point</a>.<sup id="cite_ref-Robbins_15-2" class="reference"><a href="#cite_note-Robbins-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> This is where the cell checks whether it has enough raw materials to fully replicate its DNA (nucleotide bases, DNA synthase, chromatin, etc.). An unhealthy or malnourished cell will get stuck at this checkpoint. </p><p>The G<sub>2</sub>/M checkpoint is where the cell ensures that it has enough cytoplasm and phospholipids for two daughter cells. But sometimes more importantly, it checks to see if it is the right time to replicate. There are some situations where many cells need to all replicate simultaneously (for example, a growing embryo should have a symmetric cell distribution until it reaches the mid-blastula transition). This is done by controlling the G<sub>2</sub>/M checkpoint. </p><p>The metaphase checkpoint is a fairly minor checkpoint, in that once a cell is in metaphase, it has committed to undergoing mitosis. However that's not to say it isn't important. In this checkpoint, the cell checks to ensure that the spindle has formed and that all of the chromosomes are aligned at the spindle equator before anaphase begins.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p><p>While these are the three "main" checkpoints, not all cells have to pass through each of these checkpoints in this order to replicate. Many types of cancer are caused by mutations that allow the cells to speed through the various checkpoints or even skip them altogether. Going from S to M to S phase almost consecutively. Because these cells have lost their checkpoints, any DNA mutations that may have occurred are disregarded and passed on to the daughter cells. This is one reason why cancer cells have a tendency to exponentially acquire mutations. Aside from cancer cells, many fully differentiated cell types no longer replicate so they leave the cell cycle and stay in G<sub>0</sub> until their death. Thus removing the need for cellular checkpoints. An alternative model of the cell cycle response to DNA damage has also been proposed, known as the <a href="/wiki/Postreplication_checkpoint" title="Postreplication checkpoint">postreplication checkpoint</a>. </p><p>Checkpoint regulation plays an important role in an organism's development. In sexual reproduction, when egg fertilization occurs, when the sperm binds to the egg, it releases signalling factors that notify the egg that it has been fertilized. Among other things, this induces the now fertilized oocyte to return from its previously dormant, G<sub>0</sub>, state back into the cell cycle and on to mitotic replication and division. </p><p><a href="/wiki/P53" title="P53">p53</a> plays an important role in triggering the control mechanisms at both G<sub>1</sub>/S and G<sub>2</sub>/M checkpoints. In addition to p53, checkpoint regulators are being heavily researched for their roles in cancer growth and proliferation. </p> <div class="mw-heading mw-heading2"><h2 id="Fluorescence_imaging_of_the_cell_cycle">Fluorescence imaging of the cell cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=19" title="Edit section: Fluorescence imaging of the cell cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_(FUCCI).gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/53/Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif/150px-Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif" decoding="async" width="150" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/53/Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif/225px-Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/53/Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif/300px-Far-Red_%26_Near-infrared_Fluorescent_Ubiquitination-based_Cell_Cycle_Indicator_%28FUCCI%29.gif 2x" data-file-width="320" data-file-height="320" /></a><figcaption>Fluorescent proteins visualize the cell cycle progression. IFP2.0-hGem(1/110) fluorescence is shown in green and highlights the S/G<sub>2</sub>/M phases. <a href="/wiki/SmURFP" title="SmURFP">smURFP</a>-hCdtI(30/120) fluorescence is shown in red and highlights the G<sub>0</sub>/G<sub>1</sub> phases.</figcaption></figure> <p>Pioneering work by Atsushi Miyawaki and coworkers developed the fluorescent ubiquitination-based cell cycle indicator (<a rel="nofollow" class="external text" href="http://www.conncoll.edu/ccacad/zimmer/GFP-ww/cooluses19.html">FUCCI</a>), which enables <a href="/wiki/Fluorescence" title="Fluorescence">fluorescence</a> imaging of the cell cycle. Originally, a <a href="/wiki/Green_fluorescent_protein" title="Green fluorescent protein">green fluorescent protein</a>, mAG, was fused to hGem(1/110) and an orange <a href="/wiki/Fluorescent_protein" title="Fluorescent protein">fluorescent protein</a> (mKO<sub>2</sub>) was fused to hCdt1(30/120). Note, these fusions are fragments that contain a <a href="/wiki/Nuclear_Localization_Signal" class="mw-redirect" title="Nuclear Localization Signal">nuclear localization signal</a> and <a href="/wiki/Ubiquitination" class="mw-redirect" title="Ubiquitination">ubiquitination</a> sites for <a href="/wiki/Protein_degradation" class="mw-redirect" title="Protein degradation">degradation</a>, but are not functional proteins. The <a href="/wiki/Green_fluorescent_protein" title="Green fluorescent protein">green fluorescent protein</a> is made during the S, G<sub>2</sub>, or M phase and degraded during the G<sub>0</sub> or G<sub>1</sub> phase, while the orange <a href="/wiki/Fluorescent_protein" title="Fluorescent protein">fluorescent protein</a> is made during the G<sub>0</sub> or G<sub>1</sub> phase and destroyed during the S, G<sub>2</sub>, or M phase.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> A far-red and near-infrared FUCCI was developed using a <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacteria</a>-derived <a href="/wiki/Fluorescent_protein" title="Fluorescent protein">fluorescent protein</a> (<a href="/wiki/SmURFP" title="SmURFP">smURFP</a>) and a <a href="/wiki/Phytochrome" title="Phytochrome">bacteriophytochrome</a>-derived <a href="/wiki/Fluorescent_protein" title="Fluorescent protein">fluorescent protein</a> (<a rel="nofollow" class="external text" href="http://www.nature.com/nmeth/journal/vaop/ncurrent/fig_tab/nmeth.3935_SV2.html">movie found at this link</a>).<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> Several modifications have been made to the original FUCCI system to improve its usability in several in vitro systems and model organisms. These advancements have increased the sensitivity and accuracy of cell cycle phase detection, enabling more precise assessments of cellular proliferation<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_tumor_formation">Role in tumor formation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=20" title="Edit section: Role in tumor formation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A disregulation of the cell cycle components may lead to <a href="/wiki/Tumor" class="mw-redirect" title="Tumor">tumor</a> formation.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> As mentioned above, when some genes like the cell cycle inhibitors, <a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">RB</a>, <a href="/wiki/P53" title="P53">p53</a> etc. mutate, they may cause the cell to multiply uncontrollably, forming a tumor. Although the duration of cell cycle in tumor cells is equal to or longer than that of normal cell cycle, the proportion of cells that are in active cell division (versus quiescent cells in G<sub>0</sub> phase) in tumors is much higher than that in normal tissue.<sup id="cite_ref-Baserga_1965_54-0" class="reference"><a href="#cite_note-Baserga_1965-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Thus there is a net increase in cell number as the number of cells that die by apoptosis or senescence remains the same. </p><p>The cells which are actively undergoing cell cycle are targeted in cancer therapy as the DNA is relatively exposed during cell division and hence susceptible to damage by <a href="/wiki/Chemotherapy" title="Chemotherapy">drugs</a> or <a href="/wiki/Radiotherapy" class="mw-redirect" title="Radiotherapy">radiation</a>. This fact is made use of in cancer treatment; by a process known as <a href="/wiki/Debulking" title="Debulking">debulking</a>, a significant mass of the tumor is removed which pushes a significant number of the remaining tumor cells from G<sub>0</sub> to G<sub>1</sub> phase (due to increased availability of nutrients, oxygen, growth factors etc.). Radiation or chemotherapy following the debulking procedure kills these cells which have newly entered the cell cycle.<sup id="cite_ref-Robbins_15-3" class="reference"><a href="#cite_note-Robbins-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>The fastest cycling mammalian cells in culture, crypt cells in the intestinal epithelium, have a cycle time as short as 9 to 10 hours. Stem cells in resting mouse skin may have a cycle time of more than 200 hours. Most of this difference is due to the varying length of G<sub>1</sub>, the most variable phase of the cycle. M and S do not vary much. </p><p>In general, cells are most radiosensitive in late M and G<sub>2</sub> phases and most resistant in late S phase. For cells with a longer cell cycle time and a significantly long G<sub>1</sub> phase, there is a second peak of resistance late in G<sub>1</sub>. The pattern of resistance and sensitivity correlates with the level of sulfhydryl compounds in the cell. Sulfhydryls are natural substances that protect cells from radiation damage and tend to be at their highest levels in S and at their lowest near mitosis. </p><p><a href="/wiki/Homologous_recombination" title="Homologous recombination">Homologous recombination</a> (HR) is an accurate process for <a href="/wiki/DNA_repair" title="DNA repair">repairing DNA</a> double-strand breaks. HR is nearly absent in <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a>, is most active in <a href="/wiki/S_phase" title="S phase">S phase</a>, and declines in G<sub>2</sub>/M.<sup id="cite_ref-pmid18769152_55-0" class="reference"><a href="#cite_note-pmid18769152-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Non-homologous_end_joining" title="Non-homologous end joining">Non-homologous end joining</a>, a less accurate and more mutagenic process for repairing double strand breaks, is active throughout the cell cycle. </p> <div class="mw-heading mw-heading2"><h2 id="Cell_cycle_evolution">Cell cycle evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=21" title="Edit section: Cell cycle evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Evolution_of_the_genome">Evolution of the genome</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=22" title="Edit section: Evolution of the genome"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cell cycle must duplicate all cellular constituents and equally partition them into two daughter cells. Many constituents, such as proteins and <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>, are produced continuously throughout the cell cycle (except during <a href="/wiki/Mitosis" title="Mitosis">M-phase</a>). However, the chromosomes and other associated elements like <a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">MTOCs</a>, are duplicated just once during the cell cycle. A central component of the cell cycle is its ability to coordinate the continuous and periodic duplications of different cellular elements, which evolved with the formation of the genome. </p><p>The pre-cellular environment contained functional and self-replicating <a href="/wiki/RNA" title="RNA">RNAs</a>.<sup id="cite_ref-:2_56-0" class="reference"><a href="#cite_note-:2-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> All RNA concentrations depended on the concentrations of other RNAs that might be helping or hindering the gathering of resources. In this environment, growth was simply the continuous production of RNAs. These pre-cellular structures would have had to contend with parasitic RNAs, issues of inheritance, and copy-number control of specific RNAs.<sup id="cite_ref-:2_56-1" class="reference"><a href="#cite_note-:2-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>Partitioning "genomic" RNA from "functional" RNA helped solve these problems.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> The fusion of multiple RNAs into a genome gave a template from which functional RNAs were cleaved. Now, parasitic RNAs would have to incorporate themselves into the genome, a much greater barrier, in order to survive. Controlling the copy number of genomic RNA also allowed RNA concentration to be determined through synthesis rates and RNA half-lives, instead of competition.<sup id="cite_ref-:2_56-2" class="reference"><a href="#cite_note-:2-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> Separating the duplication of genomic RNAs from the generation of functional RNAs allowed for much greater duplication fidelity of genomic RNAs without compromising the production of functional RNAs. Finally, the replacement of genomic RNA with <a href="/wiki/DNA" title="DNA">DNA</a>, which is a more stable molecule, allowed for larger genomes. The transition from self-catalysis enzyme synthesis to genome-directed enzyme synthesis was a critical step in cell evolution, and had lasting implications on the cell cycle, which must regulate functional synthesis and genomic duplication in very different ways.<sup id="cite_ref-:2_56-3" class="reference"><a href="#cite_note-:2-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cyclin-dependent_kinase_and_cyclin_evolution">Cyclin-dependent kinase and cyclin evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=23" title="Edit section: Cyclin-dependent kinase and cyclin evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cell-cycle progression is controlled by the oscillating concentrations of different <a href="/wiki/Cyclin" title="Cyclin">cyclins</a> and the resulting molecular interactions from the various <a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">cyclin-dependent kinases</a> (CDKs). In yeast, just one CDK (Cdc28 in <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">S. cerevisiae</a></i> and Cdc2 in <i><a href="/wiki/Schizosaccharomyces_pombe" title="Schizosaccharomyces pombe">S. pombe</a></i>) controls the cell cycle.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> However, in animals, whole families of CDKs have evolved.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> Cdk1 controls entry to mitosis and Cdk2, Cdk4, and Cdk6 regulate entry into S phase. Despite the evolution of the CDK family in animals, these proteins have related or redundant functions.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> For example, <i>cdk2 cdk4 cdk6</i> triple knockout mice cells can still progress through the basic cell cycle.<sup id="cite_ref-:3_65-0" class="reference"><a href="#cite_note-:3-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> <i>cdk1</i> knockouts are lethal, which suggests an ancestral CDK1-type kinase ultimately controlling the cell cycle.<sup id="cite_ref-:3_65-1" class="reference"><a href="#cite_note-:3-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> </p><p><i><a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a></i> has a Cdk1 homolog called CDKA;1, however <i>cdka;1</i> <i>A. thaliana</i> mutants are still viable,<sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> running counter to the <a href="/wiki/Opisthokont" title="Opisthokont">opisthokont</a> pattern of CDK1-type kinases as essential regulators controlling the cell cycle.<sup id="cite_ref-:4_67-0" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> Plants also have a unique group of B-type CDKs, whose functions may range from development-specific functions to major players in mitotic regulation.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="G1/S_checkpoint_evolution"><span id="G1.2FS_checkpoint_evolution"></span>G1/S checkpoint evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=24" title="Edit section: G1/S checkpoint evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:G1-S_checkpoint_regulation_across_eukaryotes.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/G1-S_checkpoint_regulation_across_eukaryotes.jpg/220px-G1-S_checkpoint_regulation_across_eukaryotes.jpg" decoding="async" width="220" height="166" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/G1-S_checkpoint_regulation_across_eukaryotes.jpg/330px-G1-S_checkpoint_regulation_across_eukaryotes.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/G1-S_checkpoint_regulation_across_eukaryotes.jpg/440px-G1-S_checkpoint_regulation_across_eukaryotes.jpg 2x" data-file-width="813" data-file-height="613" /></a><figcaption>Overviews of the G1/S transition control networks in plants, animals, and yeast. All three show striking network topology similarities, even though individual proteins in the network have very little sequence similarity.<sup id="cite_ref-:4_67-1" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The <a href="/wiki/Restriction_point" title="Restriction point">G1/S checkpoint</a> is the point at which the cell commits to division through the cell cycle. Complex regulatory networks lead to the G1/S transition decision. Across opisthokonts, there are both highly diverged protein sequences as well as strikingly similar network topologies.<sup id="cite_ref-:4_67-2" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_70-0" class="reference"><a href="#cite_note-:5-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p><p>Entry into S-phase in both yeast and animals is controlled by the levels of two opposing regulators.<sup id="cite_ref-:4_67-3" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> The networks regulating these <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> are double-negative feedback loops and positive feedback loops in both yeast and animals.<sup id="cite_ref-:4_67-4" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_70-1" class="reference"><a href="#cite_note-:5-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> Additional regulation of the regulatory network for the G1/S checkpoint in yeast and animals includes the <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a>/de-phosphorylation of CDK-cyclin complexes. The sum of these regulatory networks creates a <a href="/wiki/Hysteresis" title="Hysteresis">hysteretic</a> and bistable scheme, despite the specific proteins being highly diverged.<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> For yeast, <a href="/wiki/Whi5" title="Whi5">Whi5</a> must be suppressed by Cln3 phosphorylation for SBF to be expressed,<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> while in animals <a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">Rb</a> must be suppressed by the Cdk4/6-cyclin D complex for <a href="/wiki/E2F" title="E2F">E2F</a> to be expressed.<sup id="cite_ref-:6_75-0" class="reference"><a href="#cite_note-:6-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> Both Rb and Whi5 inhibit transcript through the recruitment of histone deacetylase proteins to promoters.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> Both proteins additionally have multiple CDK phosphorylation sites through which they are inhibited.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:6_75-1" class="reference"><a href="#cite_note-:6-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> However, these proteins share no sequence similarity. </p><p>Studies in <i>A. thaliana</i> extend our knowledge of the G1/S transition across <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> as a whole. Plants also share a number of conserved network features with opisthokonts, and many plant regulators have direct animal homologs.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> For example, plants also need to suppress Rb for E2F translation in the network.<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> These conserved elements of the plant and animal cell cycles may be ancestral in eukaryotes. While yeast share a conserved network topology with plants and animals, the highly diverged nature of yeast regulators suggests possible rapid evolution along the yeast lineage.<sup id="cite_ref-:4_67-5" class="reference"><a href="#cite_note-:4-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=25" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Cellular_model" title="Cellular model">Cellular model</a></li> <li><a href="/wiki/Eukaryotic_DNA_replication" title="Eukaryotic DNA replication">Eukaryotic DNA replication</a></li> <li><a href="/wiki/Mitotic_catastrophe" title="Mitotic catastrophe">Mitotic catastrophe</a></li> <li><a href="/wiki/Origin_recognition_complex" title="Origin recognition complex">Origin recognition complex</a></li> <li><a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">Retinoblastoma protein</a></li> <li><a href="/wiki/Synchronous_culture" title="Synchronous culture">Synchronous culture</a> – synchronization of cell cultures</li> <li><a href="/wiki/Wee1" title="Wee1">Wee1</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=Cell_cycle&action=edit&section=26" 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: 33em;"> <ol class="references"> <li id="cite_note-Alberts2019-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Alberts2019_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFAlbertsHopkinJohnsonMorgan2019" class="citation book cs1">Alberts B, Hopkin K, Johnson A, Morgan D, Raff M, Roberts K, Walter P (2019). <i>Essential cell biology</i> (Fifth ed.). 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Palanivelu R (ed.). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3410867">"A general G1/S-phase cell-cycle control module in the flowering plant Arabidopsis thaliana"</a>. <i>PLOS Genetics</i>. <b>8</b> (8): e1002847. <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/10.1371%2Fjournal.pgen.1002847">10.1371/journal.pgen.1002847</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/PMC3410867">3410867</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/22879821">22879821</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+Genetics&rft.atitle=A+general+G1%2FS-phase+cell-cycle+control+module+in+the+flowering+plant+Arabidopsis+thaliana&rft.volume=8&rft.issue=8&rft.pages=e1002847&rft.date=2012-08-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3410867%23id-name%3DPMC&rft_id=info%3Apmid%2F22879821&rft_id=info%3Adoi%2F10.1371%2Fjournal.pgen.1002847&rft.aulast=Zhao&rft.aufirst=X&rft.au=Harashima%2C+H&rft.au=Dissmeyer%2C+N&rft.au=Pusch%2C+S&rft.au=Weimer%2C+AK&rft.au=Bramsiepe%2C+J&rft.au=Bouyer%2C+D&rft.au=Rademacher%2C+S&rft.au=Nowack%2C+MK&rft.au=Novak%2C+B&rft.au=Sprunck%2C+S&rft.au=Schnittger%2C+A&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3410867&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></span> </li> <li id="cite_note-80"><span class="mw-cite-backlink"><b><a href="#cite_ref-80">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWeimerNowackBouyerZhao2012" class="citation journal cs1">Weimer AK, Nowack MK, Bouyer D, Zhao X, Harashima H, Naseer S, et al. (October 2012). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3517237">"Retinoblastoma related1 regulates asymmetric cell divisions in Arabidopsis"</a>. <i>The Plant Cell</i>. <b>24</b> (10): <span class="nowrap">4083–</span>4095. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1105%2Ftpc.112.104620">10.1105/tpc.112.104620</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/PMC3517237">3517237</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/23104828">23104828</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Plant+Cell&rft.atitle=Retinoblastoma+related1+regulates+asymmetric+cell+divisions+in+Arabidopsis&rft.volume=24&rft.issue=10&rft.pages=%3Cspan+class%3D%22nowrap%22%3E4083-%3C%2Fspan%3E4095&rft.date=2012-10&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3517237%23id-name%3DPMC&rft_id=info%3Apmid%2F23104828&rft_id=info%3Adoi%2F10.1105%2Ftpc.112.104620&rft.aulast=Weimer&rft.aufirst=AK&rft.au=Nowack%2C+MK&rft.au=Bouyer%2C+D&rft.au=Zhao%2C+X&rft.au=Harashima%2C+H&rft.au=Naseer%2C+S&rft.au=De+Winter%2C+F&rft.au=Dissmeyer%2C+N&rft.au=Geldner%2C+N&rft.au=Schnittger%2C+A&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3517237&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cell_cycle&action=edit&section=27" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 33em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFMorgan2007" class="citation book cs1">Morgan DO (2007). <i>The Cell Cycle: Principles of Control</i>. London: Published by New Science Press in association with Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87893-508-6" title="Special:BookSources/978-0-87893-508-6"><bdi>978-0-87893-508-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Cell+Cycle%3A+Principles+of+Control&rft.place=London&rft.pub=Published+by+New+Science+Press+in+association+with+Oxford+University+Press&rft.date=2007&rft.isbn=978-0-87893-508-6&rft.aulast=Morgan&rft.aufirst=DO&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAlbertsJohnsonLewisRaff2008" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2008). "Chapter 17". <i>Molecular Biology of the Cell</i> (5th ed.). New York: Garland Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-4111-6" title="Special:BookSources/978-0-8153-4111-6"><bdi>978-0-8153-4111-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+17&rft.btitle=Molecular+Biology+of+the+Cell&rft.place=New+York&rft.edition=5th&rft.pub=Garland+Science&rft.date=2008&rft.isbn=978-0-8153-4111-6&rft.aulast=Alberts&rft.aufirst=B&rft.au=Johnson%2C+A&rft.au=Lewis%2C+J&rft.au=Raff%2C+M&rft.au=Roberts%2C+K&rft.au=Walter%2C+P&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFKriegerScottMatsudairaLodish2004" class="citation book cs1">Krieger M, Scott MP, Matsudaira PT, Lodish HF, Darnell JE, Zipursky L, Kaiser C, Berk A (2004). <a rel="nofollow" class="external text" href="https://archive.org/details/molecularcellbio00harv"><i>Molecular cell biology</i></a>. New York: W.H. Freeman and CO. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7167-4366-8" title="Special:BookSources/978-0-7167-4366-8"><bdi>978-0-7167-4366-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+cell+biology&rft.place=New+York&rft.pub=W.H.+Freeman+and+CO&rft.date=2004&rft.isbn=978-0-7167-4366-8&rft.aulast=Krieger&rft.aufirst=M&rft.au=Scott%2C+MP&rft.au=Matsudaira%2C+PT&rft.au=Lodish%2C+HF&rft.au=Darnell%2C+JE&rft.au=Zipursky%2C+L&rft.au=Kaiser%2C+C&rft.au=Berk%2C+A&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmolecularcellbio00harv&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWatsonBakerBellGann2004" class="citation book cs1">Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R (2004). "Chapter 7". <i>Molecular biology of the gene</i> (5th ed.). San Francisco: Pearson/Benjamin Cummings. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8053-4642-8" title="Special:BookSources/978-0-8053-4642-8"><bdi>978-0-8053-4642-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+7&rft.btitle=Molecular+biology+of+the+gene&rft.place=San+Francisco&rft.edition=5th&rft.pub=Pearson%2FBenjamin+Cummings&rft.date=2004&rft.isbn=978-0-8053-4642-8&rft.aulast=Watson&rft.aufirst=JD&rft.au=Baker%2C+TA&rft.au=Bell%2C+SP&rft.au=Gann%2C+A&rft.au=Levine%2C+M&rft.au=Losick%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></li></ul> </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=Cell_cycle&action=edit&section=28" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><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/40px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/60px-Commons-logo.svg.png 1.5x" data-file-width="1024" data-file-height="1376" /></a></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Cell_cycle" class="extiw" title="commons:Category:Cell cycle">Cell cycle</a></span>.</div></div> </div> <ul><li><span class="noviewer" typeof="mw:File"><span><img alt="Public Domain" src="//upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/12px-PD-icon.svg.png" decoding="async" width="12" height="12" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/18px-PD-icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/24px-PD-icon.svg.png 2x" data-file-width="196" data-file-height="196" /></span></span> This article incorporates <a href="/wiki/Copyright_status_of_works_by_the_federal_government_of_the_United_States" title="Copyright status of works by the federal government of the United States">public domain material</a> from <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite class="citation cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20091208004309/https://www.ncbi.nlm.nih.gov/About/primer/index.html"><i>Science Primer</i></a>. <a href="/wiki/NCBI" class="mw-redirect" title="NCBI">NCBI</a>. Archived from <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/About/primer/index.html">the original</a> on 8 December 2009.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Science+Primer&rft.pub=NCBI&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2FAbout%2Fprimer%2Findex.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+cycle" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://www.ibiology.org/cell-biology/controlling-cell-cycle/">David Morgan's Seminar: Controlling the Cell Cycle</a></li> <li><a rel="nofollow" class="external text" href="https://www.kasem.info/histology-lectures/first-year-histology-lectures/the-cell-cycle">The cell cycle & Cell death</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20181030131240/https://www.kasem.info/histology-lectures/first-year-histology-lectures/the-cell-cycle">Archived</a> 30 October 2018 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="http://www.cellcycle.info">Transcriptional program of the cell cycle: high-resolution timing</a></li> <li><a rel="nofollow" class="external text" href="http://www.sceptrans.org">Cell cycle and metabolic cycle regulated transcription in yeast</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20160606092520/http://1lec.com/control-of-cell-cycle/">Cell Cycle Animation</a> 1Lec.com</li> <li><a rel="nofollow" class="external text" href="http://www.landesbioscience.com/journals/cc/index.php">Cell Cycle</a></li> <li><a rel="nofollow" class="external text" href="http://www.conncoll.edu/ccacad/zimmer/GFP-ww/cooluses19.html">Fucci:Using GFP to visualize the cell-cycle</a></li> <li><a rel="nofollow" class="external text" href="http://www.scq.ubc.ca/?p=248">Science Creative Quarterly's overview of the cell cycle</a></li> <li><a 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a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cell_cycle_proteins" title="Template:Cell cycle proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cell_cycle_proteins" title="Template talk:Cell cycle proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cell_cycle_proteins" title="Special:EditPage/Template:Cell cycle proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_cycle_proteins179" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Cell cycle</a> <a href="/wiki/Protein" 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 class="mw-selflink-fragment" href="#Inhibitors">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 href="/wiki/Anaphase" title="Anaphase">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 class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" 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