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class="vector-toc-numb">2</span> <span>Activation mechanisms</span> </div> </a> <button aria-controls="toc-Activation_mechanisms-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 Activation mechanisms subsection</span> </button> <ul id="toc-Activation_mechanisms-sublist" class="vector-toc-list"> <li id="toc-Intrinsic_pathway" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Intrinsic_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Intrinsic pathway</span> </div> </a> <ul id="toc-Intrinsic_pathway-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Extrinsic_pathway" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Extrinsic_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Extrinsic pathway</span> </div> </a> <ul id="toc-Extrinsic_pathway-sublist" class="vector-toc-list"> <li id="toc-TNF_pathway" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#TNF_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.1</span> <span>TNF pathway</span> </div> </a> <ul id="toc-TNF_pathway-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Fas_pathway" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Fas_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.2</span> <span>Fas pathway</span> </div> </a> <ul id="toc-Fas_pathway-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Common_components" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Common_components"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.3</span> <span>Common components</span> </div> </a> <ul id="toc-Common_components-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Caspases" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Caspases"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.4</span> <span>Caspases</span> </div> </a> <ul id="toc-Caspases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Caspase-independent_apoptotic_pathway" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Caspase-independent_apoptotic_pathway"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.5</span> <span>Caspase-independent apoptotic pathway</span> </div> </a> <ul id="toc-Caspase-independent_apoptotic_pathway-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Apoptosis_model_in_amphibians" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Apoptosis_model_in_amphibians"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Apoptosis model in amphibians</span> </div> </a> <ul id="toc-Apoptosis_model_in_amphibians-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Negative_regulators_of_apoptosis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Negative_regulators_of_apoptosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Negative regulators of apoptosis</span> </div> </a> <ul id="toc-Negative_regulators_of_apoptosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Proteolytic_caspase_cascade:_Killing_the_cell" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Proteolytic_caspase_cascade:_Killing_the_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Proteolytic caspase cascade: Killing the cell</span> </div> </a> <button aria-controls="toc-Proteolytic_caspase_cascade:_Killing_the_cell-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 Proteolytic caspase cascade: Killing the cell subsection</span> </button> <ul id="toc-Proteolytic_caspase_cascade:_Killing_the_cell-sublist" class="vector-toc-list"> <li id="toc-Apoptotic_cell_disassembly" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Apoptotic_cell_disassembly"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Apoptotic cell disassembly</span> </div> </a> <ul id="toc-Apoptotic_cell_disassembly-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Removal_of_dead_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Removal_of_dead_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Removal of dead cells</span> </div> </a> <ul id="toc-Removal_of_dead_cells-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Pathway_knock-outs" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Pathway_knock-outs"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Pathway knock-outs</span> </div> </a> <ul id="toc-Pathway_knock-outs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Methods_for_distinguishing_apoptotic_from_necrotic_cells" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Methods_for_distinguishing_apoptotic_from_necrotic_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Methods for distinguishing apoptotic from necrotic cells</span> </div> </a> <ul id="toc-Methods_for_distinguishing_apoptotic_from_necrotic_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Implication_in_disease" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Implication_in_disease"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Implication in disease</span> </div> </a> <button aria-controls="toc-Implication_in_disease-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 Implication in disease subsection</span> </button> <ul id="toc-Implication_in_disease-sublist" class="vector-toc-list"> <li id="toc-Defective_pathways" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Defective_pathways"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Defective pathways</span> </div> </a> <ul id="toc-Defective_pathways-sublist" class="vector-toc-list"> <li id="toc-Dysregulation_of_p53" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Dysregulation_of_p53"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.1</span> <span>Dysregulation of p53</span> </div> </a> <ul id="toc-Dysregulation_of_p53-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Inhibition" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inhibition"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Inhibition</span> </div> </a> <ul id="toc-Inhibition-sublist" class="vector-toc-list"> <li id="toc-HeLa_cell" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#HeLa_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2.1</span> <span>HeLa cell</span> </div> </a> <ul id="toc-HeLa_cell-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Treatments" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Treatments"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2.2</span> <span>Treatments</span> </div> </a> <ul id="toc-Treatments-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Hyperactive_apoptosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Hyperactive_apoptosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Hyperactive apoptosis</span> </div> </a> <ul id="toc-Hyperactive_apoptosis-sublist" class="vector-toc-list"> <li id="toc-Treatments_2" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Treatments_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.1</span> <span>Treatments</span> </div> </a> <ul id="toc-Treatments_2-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-HIV_progression" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#HIV_progression"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>HIV progression</span> </div> </a> <ul id="toc-HIV_progression-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Viral_infection" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Viral_infection"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.5</span> <span>Viral infection</span> </div> </a> <ul id="toc-Viral_infection-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Plants" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Plants"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Plants</span> </div> </a> <ul id="toc-Plants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Caspase-independent_apoptosis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Caspase-independent_apoptosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Caspase-independent apoptosis</span> </div> </a> <ul id="toc-Caspase-independent_apoptosis-sublist" class="vector-toc-list"> </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">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Explanatory_footnotes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Explanatory_footnotes"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Explanatory footnotes</span> </div> </a> <ul id="toc-Explanatory_footnotes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Citations" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Citations"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>Citations</span> </div> </a> <ul id="toc-Citations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-General_bibliography" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#General_bibliography"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>General bibliography</span> </div> </a> <ul id="toc-General_bibliography-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">14</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Apoptosis</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 63 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-63" 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">63 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Apoptose" title="Apoptose – Afrikaans" lang="af" hreflang="af" data-title="Apoptose" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A7%D8%B3%D8%AA%D9%85%D8%A7%D8%AA%D8%A9_(%D8%B9%D9%84%D9%85_%D8%A3%D8%AD%D9%8A%D8%A7%D8%A1)" title="استماتة (علم أحياء) – Arabic" lang="ar" hreflang="ar" data-title="استماتة (علم أحياء)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Apoptoz" title="Apoptoz – Azerbaijani" lang="az" hreflang="az" data-title="Apoptoz" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%95%E0%A7%8B%E0%A6%B7%E0%A6%AA%E0%A6%A4%E0%A6%A8" title="কোষপতন – Bangla" lang="bn" hreflang="bn" data-title="কোষপতন" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%90%D0%BF%D0%B0%D0%BF%D1%82%D0%BE%D0%B7" title="Апаптоз – Belarusian" lang="be" hreflang="be" data-title="Апаптоз" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%90%D0%BF%D0%B0%D0%BF%D1%82%D0%BE%D0%B7" title="Апаптоз – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Апаптоз" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%90%D0%BF%D0%BE%D0%BF%D1%82%D0%BE%D0%B7%D0%B0" title="Апоптоза – Bulgarian" lang="bg" hreflang="bg" data-title="Апоптоза" 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/Apoptoza" title="Apoptoza – Bosnian" lang="bs" hreflang="bs" data-title="Apoptoza" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Apoptosi" title="Apoptosi – Catalan" lang="ca" hreflang="ca" data-title="Apoptosi" 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/Apopt%C3%B3za" title="Apoptóza – Czech" lang="cs" hreflang="cs" data-title="Apoptóza" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Apoptose" title="Apoptose – Danish" lang="da" hreflang="da" data-title="Apoptose" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-ary mw-list-item"><a href="https://ary.wikipedia.org/wiki/%D9%85%D9%88%D8%AA_%D8%AE%D9%84%D9%88%D9%8A_%D9%85%D8%A8%D8%B1%D9%85%D8%AC" title="موت خلوي مبرمج – Moroccan Arabic" lang="ary" hreflang="ary" data-title="موت خلوي مبرمج" data-language-autonym="الدارجة" data-language-local-name="Moroccan Arabic" class="interlanguage-link-target"><span>الدارجة</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Apoptose" title="Apoptose – German" lang="de" hreflang="de" data-title="Apoptose" 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/Apoptoos" title="Apoptoos – Estonian" lang="et" hreflang="et" data-title="Apoptoos" 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%91%CF%80%CF%8C%CF%80%CF%84%CF%89%CF%83%CE%B7" title="Απόπτωση – Greek" lang="el" hreflang="el" data-title="Απόπτωση" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Spanish" lang="es" hreflang="es" data-title="Apoptosis" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-ext mw-list-item"><a href="https://ext.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Extremaduran" lang="ext" hreflang="ext" data-title="Apoptosis" data-language-autonym="Estremeñu" data-language-local-name="Extremaduran" class="interlanguage-link-target"><span>Estremeñu</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Apoptosi" title="Apoptosi – Basque" lang="eu" hreflang="eu" data-title="Apoptosi" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A2%D9%BE%D9%88%D9%BE%D8%AA%D9%88%D8%B2" title="آپوپتوز – Persian" lang="fa" hreflang="fa" data-title="آپوپتوز" 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/Apoptose" title="Apoptose – French" lang="fr" hreflang="fr" data-title="Apoptose" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Apapt%C3%B3is" title="Apaptóis – Irish" lang="ga" hreflang="ga" data-title="Apaptóis" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Apoptose" title="Apoptose – Galician" lang="gl" hreflang="gl" data-title="Apoptose" 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%9E%90%EC%82%B4" title="세포자살 – Korean" lang="ko" hreflang="ko" data-title="세포자살" 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%B1%D5%BA%D5%B8%D5%BA%D5%BF%D5%B8%D5%A6" title="Ապոպտոզ – Armenian" lang="hy" hreflang="hy" data-title="Ապոպտոզ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Apoptozo" title="Apoptozo – Ido" lang="io" hreflang="io" data-title="Apoptozo" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Indonesian" lang="id" hreflang="id" data-title="Apoptosis" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Apoptosi" title="Apoptosi – Italian" lang="it" hreflang="it" data-title="Apoptosi" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%90%D7%A4%D7%95%D7%A4%D7%98%D7%95%D7%96%D7%94" title="אפופטוזה – Hebrew" lang="he" hreflang="he" data-title="אפופטוזה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%85%E0%B2%AA%E0%B3%8A%E0%B2%AA%E0%B3%8D%E0%B2%9F%E0%B3%8B%E0%B2%B8%E0%B2%BF%E0%B2%B8%E0%B3%8D" title="ಅಪೊಪ್ಟೋಸಿಸ್ – Kannada" lang="kn" hreflang="kn" data-title="ಅಪೊಪ್ಟೋಸಿಸ್" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%90%E1%83%9E%E1%83%9D%E1%83%9E%E1%83%A2%E1%83%9D%E1%83%96%E1%83%98" title="აპოპტოზი – Georgian" lang="ka" hreflang="ka" data-title="აპოპტოზი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%90%D0%BF%D0%BE%D0%BF%D0%B5%D0%BE%D0%B7_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F)" title="Апопеоз (биология) – Kyrgyz" lang="ky" hreflang="ky" data-title="Апопеоз (биология)" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Latin" lang="la" hreflang="la" data-title="Apoptosis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Apoptoz%C4%97" title="Apoptozė – Lithuanian" lang="lt" hreflang="lt" data-title="Apoptozė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Apopt%C3%B3zis" title="Apoptózis – Hungarian" lang="hu" hreflang="hu" data-title="Apoptózis" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%90%D0%BF%D0%BE%D0%BF%D1%82%D0%BE%D0%B7%D0%B0" title="Апоптоза – Macedonian" lang="mk" hreflang="mk" data-title="Апоптоза" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%95%E0%B5%8B%E0%B4%B6%E0%B4%BE%E0%B4%A4%E0%B5%8D%E0%B4%AE%E0%B4%B9%E0%B4%A4%E0%B5%8D%E0%B4%AF" title="കോശാത്മഹത്യ – Malayalam" lang="ml" hreflang="ml" data-title="കോശാത്മഹത്യ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Apoptose" title="Apoptose – Dutch" lang="nl" hreflang="nl" data-title="Apoptose" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%A2%E3%83%9D%E3%83%88%E3%83%BC%E3%82%B7%E3%82%B9" title="アポトーシス – Japanese" lang="ja" hreflang="ja" data-title="アポトーシス" 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/Apoptose" title="Apoptose – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Apoptose" 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/Apoptose" title="Apoptose – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Apoptose" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Apopt%C3%B2si" title="Apoptòsi – Occitan" lang="oc" hreflang="oc" data-title="Apoptòsi" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Apoptoz" title="Apoptoz – Uzbek" lang="uz" hreflang="uz" data-title="Apoptoz" 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/Apoptoza" title="Apoptoza – Polish" lang="pl" hreflang="pl" data-title="Apoptoza" 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/Apoptose" title="Apoptose – Portuguese" lang="pt" hreflang="pt" data-title="Apoptose" 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/Apoptoz%C4%83" title="Apoptoză – Romanian" lang="ro" hreflang="ro" data-title="Apoptoză" 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%90%D0%BF%D0%BE%D0%BF%D1%82%D0%BE%D0%B7" title="Апоптоз – Russian" lang="ru" hreflang="ru" data-title="Апоптоз" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Simple English" lang="en-simple" hreflang="en-simple" data-title="Apoptosis" 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/Apopt%C3%B3za" title="Apoptóza – Slovak" lang="sk" hreflang="sk" data-title="Apoptóza" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://sl.wikipedia.org/wiki/Apoptoza" title="Apoptoza – Slovenian" lang="sl" hreflang="sl" data-title="Apoptoza" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%90%D0%BF%D0%BE%D0%BF%D1%82%D0%BE%D0%B7%D0%B0" title="Апоптоза – Serbian" lang="sr" hreflang="sr" data-title="Апоптоза" 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/Apoptoza" title="Apoptoza – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Apoptoza" 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-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Apoptosis" title="Apoptosis – Sundanese" lang="su" hreflang="su" data-title="Apoptosis" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Apoptoosi" title="Apoptoosi – Finnish" lang="fi" hreflang="fi" data-title="Apoptoosi" 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/Apoptos" title="Apoptos – Swedish" lang="sv" hreflang="sv" data-title="Apoptos" 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/Apoptosis" title="Apoptosis – Tagalog" lang="tl" hreflang="tl" data-title="Apoptosis" 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%89%E0%AE%AF%E0%AE%BF%E0%AE%B0%E0%AE%A3%E0%AF%81_%E0%AE%A4%E0%AE%A9%E0%AF%8D%E0%AE%AE%E0%AE%9F%E0%AE%BF%E0%AE%B5%E0%AF%81" title="உயிரணு தன்மடிவு – Tamil" lang="ta" hreflang="ta" data-title="உயிரணு தன்மடிவு" 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%AD%E0%B8%B0%E0%B8%9E%E0%B8%AD%E0%B8%9E%E0%B9%82%E0%B8%97%E0%B8%8B%E0%B8%B4%E0%B8%AA" title="อะพอพโทซิส – Thai" lang="th" hreflang="th" data-title="อะพอพโทซิส" 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/Apoptoz" title="Apoptoz – Turkish" lang="tr" hreflang="tr" data-title="Apoptoz" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%90%D0%BF%D0%BE%D0%BF%D1%82%D0%BE%D0%B7" title="Апоптоз – Ukrainian" lang="uk" hreflang="uk" data-title="Апоптоз" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%A7%D8%B3%D8%AA%D9%85%D8%A7%D8%AA%D8%AA" title="استماتت – Urdu" lang="ur" hreflang="ur" data-title="استماتت" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Ch%E1%BA%BFt_t%E1%BA%BF_b%C3%A0o_theo_ch%C6%B0%C6%A1ng_tr%C3%ACnh" title="Chết tế bào theo chương trình – Vietnamese" lang="vi" hreflang="vi" data-title="Chết tế bào theo chương trình" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E7%BB%86%E8%83%9E%E5%87%8B%E4%BA%A1" title="细胞凋亡 – Wu" lang="wuu" hreflang="wuu" data-title="细胞凋亡" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%BB%86%E8%83%9E%E5%87%8B%E4%BA%A1" title="细胞凋亡 – Chinese" lang="zh" hreflang="zh" data-title="细胞凋亡" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q14599311#sitelinks-wikipedia" title="Edit 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searchaux" style="display:none">Type of programmed cell death in multicellular organisms</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:dimgray; color: white">Apoptosis</th></tr><tr><td colspan="2" class="infobox-image"><span typeof="mw:File"><a href="/wiki/File:Apoptosis_DU145_cells_mosaic.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Apoptosis_DU145_cells_mosaic.jpg/306px-Apoptosis_DU145_cells_mosaic.jpg" decoding="async" width="306" height="614" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Apoptosis_DU145_cells_mosaic.jpg/459px-Apoptosis_DU145_cells_mosaic.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/2/23/Apoptosis_DU145_cells_mosaic.jpg 2x" data-file-width="503" data-file-height="1009" /></a></span><div class="infobox-caption">An <a href="/wiki/Etoposide" title="Etoposide">etoposide</a>-treated <a href="/wiki/DU145" title="DU145">DU145 prostate cancer cell</a> exploding into a cascade of apoptotic bodies. The sub images were extracted from a 61-hour <a href="/wiki/Time-lapse_microscopy" title="Time-lapse microscopy">time-lapse microscopy</a> video, created using <a href="/wiki/Quantitative_phase-contrast_microscopy" title="Quantitative phase-contrast microscopy">quantitative phase-contrast microscopy</a>. The optical thickness is color-coded. With increasing thickness, color changes from gray to yellow, red, purple and finally black. <a rel="nofollow" class="external text" href="http://www.cellimagelibrary.org/images/43705"><br />See the video at The Cell: An Image Library</a></div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D017209">D017209</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terminology" title="Anatomical terminology">Anatomical terminology</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q14599311" class="extiw" title="d:Q14599311">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p><b>Apoptosis</b> (from <a href="/wiki/Ancient_Greek_language" class="mw-redirect" title="Ancient Greek language">Ancient Greek</a>: <span lang="grc"><a href="https://en.wiktionary.org/wiki/%E1%BC%80%CF%80%CF%8C%CF%80%CF%84%CF%89%CF%83%CE%B9%CF%82" class="extiw" title="wikt:ἀπόπτωσις">ἀπόπτωσις</a></span>, <small><a href="/wiki/Romanization_of_Ancient_Greek" class="mw-redirect" title="Romanization of Ancient Greek">romanized</a>: </small><span title="Ancient Greek-language romanization"><i lang="grc-Latn">apóptōsis</i></span>, <small><a href="/wiki/Literal_translation" title="Literal translation">lit.</a> </small>'falling off') is a form of <a href="/wiki/Programmed_cell_death" title="Programmed cell death">programmed cell death</a> that occurs in <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellular organisms</a> and in some eukaryotic, single-celled microorganisms such as <a href="/wiki/Yeast" title="Yeast">yeast</a>.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Biochemistry" title="Biochemistry">Biochemical</a> events lead to characteristic cell changes (<a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>) and death.<sup id="cite_ref-pmid14499155_2-0" class="reference"><a href="#cite_note-pmid14499155-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These changes include <a href="/wiki/Bleb_(cell_biology)" title="Bleb (cell biology)">blebbing</a>, <a href="/wiki/Plasmolysis" title="Plasmolysis">cell shrinkage</a>, <a href="/wiki/Karyorrhexis" title="Karyorrhexis">nuclear fragmentation</a>, <a href="/wiki/Pyknosis" title="Pyknosis">chromatin condensation</a>, <a href="/wiki/Apoptotic_DNA_fragmentation" title="Apoptotic DNA fragmentation">DNA fragmentation</a>, and <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> decay. The average adult human loses 50 to 70 <a href="/wiki/1,000,000,000" title="1,000,000,000">billion</a> cells each day due to apoptosis.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> For the average human child between 8 and 14 years old, each day the approximate loss is 20 to 30 billion cells.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>In contrast to <a href="/wiki/Necrosis" title="Necrosis">necrosis</a>, which is a form of traumatic cell death that results from acute cellular injury, apoptosis is a highly regulated and controlled process that confers advantages during an organism's life cycle. For example, the separation of fingers and toes in a developing human <a href="/wiki/Embryo" title="Embryo">embryo</a> occurs because cells between the digits undergo apoptosis. Unlike necrosis, apoptosis produces cell fragments called <a href="/wiki/Extracellular_vesicle#Apoptotic_bodies" title="Extracellular vesicle">apoptotic bodies</a> that <a href="/wiki/Phagocyte" title="Phagocyte">phagocytes</a> are able to engulf and remove before the contents of the cell can spill out onto surrounding cells and cause damage to them.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Because apoptosis cannot stop once it has begun, it is a highly regulated process. Apoptosis can be initiated through one of two pathways. In the <i>intrinsic pathway</i> the cell kills itself because it senses <a href="/wiki/Cellular_stress_response" title="Cellular stress response">cell stress</a>, while in the <i>extrinsic pathway</i> the cell kills itself because of signals from other cells. Weak external signals may also activate the intrinsic pathway of apoptosis.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Both pathways induce cell death by activating <a href="/wiki/Caspase" title="Caspase">caspases</a>, which are <a href="/wiki/Protease" title="Protease">proteases</a>, or enzymes that degrade proteins. The two pathways both activate initiator caspases, which then activate executioner caspases, which then kill the cell by degrading proteins indiscriminately. </p><p>In addition to its importance as a biological phenomenon, defective apoptotic processes have been implicated in a wide variety of diseases. Excessive apoptosis causes <a href="/wiki/Atrophy" title="Atrophy">atrophy</a>, whereas an insufficient amount results in uncontrolled cell proliferation, such as <a href="/wiki/Cancer" title="Cancer">cancer</a>. Some factors like <a href="/wiki/Fas_receptor" title="Fas receptor">Fas receptors</a> and caspases promote apoptosis, while some members of the <a href="/wiki/Bcl-2_family" title="Bcl-2 family">Bcl-2 family</a> of proteins inhibit apoptosis.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Discovery_and_etymology">Discovery and etymology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=1" title="Edit section: Discovery and etymology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/History_of_apoptosis_research" title="History of apoptosis research">History of apoptosis research</a></div> <p>German scientist <a href="/wiki/Carl_Vogt" title="Carl Vogt">Carl Vogt</a> was first to describe the principle of apoptosis in 1842. In 1885, anatomist <a href="/wiki/Walther_Flemming" title="Walther Flemming">Walther Flemming</a> delivered a more precise description of the process of programmed cell death. However, it was not until 1965 that the topic was resurrected. While studying tissues using electron microscopy, <a href="/wiki/John_Kerr_(pathologist)" title="John Kerr (pathologist)">John Kerr</a> at the University of Queensland was able to distinguish apoptosis from traumatic cell death.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Following the publication of a paper describing the phenomenon, Kerr was invited to join <a href="/wiki/Alastair_Currie" title="Alastair Currie">Alastair Currie</a>, as well as <a href="/wiki/Andrew_Wyllie_(pathologist)" title="Andrew Wyllie (pathologist)">Andrew Wyllie</a>, who was Currie's graduate student,<sup id="cite_ref-AHW_1972_10-0" class="reference"><a href="#cite_note-AHW_1972-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> at the <a href="/wiki/University_of_Aberdeen" title="University of Aberdeen">University of Aberdeen</a>. In 1972, the trio published a seminal article in the <i><a href="/wiki/British_Journal_of_Cancer" title="British Journal of Cancer">British Journal of Cancer</a></i>.<sup id="cite_ref-Kerr1972_11-0" class="reference"><a href="#cite_note-Kerr1972-11"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Kerr had initially used the term programmed cell necrosis, but in the article, the process of natural cell death was called <i>apoptosis</i>. Kerr, Wyllie and Currie credited James Cormack, a professor of Greek language at University of Aberdeen, with suggesting the term apoptosis. Kerr received the <a href="/wiki/Paul_Ehrlich_and_Ludwig_Darmstaedter_Prize" title="Paul Ehrlich and Ludwig Darmstaedter Prize">Paul Ehrlich and Ludwig Darmstaedter Prize</a> on March 14, 2000, for his description of apoptosis. He shared the prize with Boston biologist <a href="/wiki/H._Robert_Horvitz" title="H. Robert Horvitz">H. Robert Horvitz</a>.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>For many years, neither "apoptosis" nor "programmed cell death" was a highly cited term. Two discoveries brought cell death from obscurity to a major field of research: identification of the first component of the cell death control and effector mechanisms, and linkage of abnormalities in cell death to human disease, in particular cancer. This occurred in 1988 when it was shown that BCL2, the gene responsible for follicular lymphoma, encoded a protein that inhibited cell death.<sup id="cite_ref-bcl2_13-0" class="reference"><a href="#cite_note-bcl2-13"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>The 2002 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Medicine</a> was awarded to <a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Sydney Brenner</a>, <a href="/wiki/H._Robert_Horvitz" title="H. Robert Horvitz">H. Robert Horvitz</a> and <a href="/wiki/John_Sulston" title="John Sulston">John Sulston</a> for their work identifying genes that control apoptosis. The genes were identified by studies in the nematode <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i> and homologues of these genes function in humans to regulate apoptosis.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:John_Sulston.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bc/John_Sulston.jpg/170px-John_Sulston.jpg" decoding="async" width="170" height="167" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bc/John_Sulston.jpg/255px-John_Sulston.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bc/John_Sulston.jpg/340px-John_Sulston.jpg 2x" data-file-width="1102" data-file-height="1083" /></a><figcaption><a href="/wiki/John_Sulston" title="John Sulston">John Sulston</a> won the <a href="/wiki/Nobel_Prize_in_Medicine" class="mw-redirect" title="Nobel Prize in Medicine">Nobel Prize in Medicine</a> in 2002, for his pioneering research on apoptosis.</figcaption></figure> <p>In Greek, apoptosis translates to the "falling off" of leaves from a tree.<sup id="cite_ref-FOOTNOTEAlberts1021_14-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts1021-14"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Cormack, professor of Greek language, reintroduced the term for medical use as it had a medical meaning for the Greeks over two thousand years before. <a href="/wiki/Hippocrates" title="Hippocrates">Hippocrates</a> used the term to mean "the falling off of the bones". <a href="/wiki/Galen" title="Galen">Galen</a> extended its meaning to "the dropping of the scabs". Cormack was no doubt aware of this usage when he suggested the name. Debate continues over the correct pronunciation, with opinion divided between a pronunciation with the second <i>p</i> silent (<span class="rt-commentedText nowrap"><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="/æ/: 'a' in 'bad'">æ</span><span title="'p' in 'pie'">p</span><span title="/ə/: 'a' in 'about'">ə</span><span title="/ˈ/: primary stress follows">ˈ</span><span title="'t' in 'tie'">t</span><span title="/oʊ/: 'o' in 'code'">oʊ</span><span title="'s' in 'sigh'">s</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="'s' in 'sigh'">s</span></span>/</a></span></span> <a href="/wiki/Help:Pronunciation_respelling_key" title="Help:Pronunciation respelling key"><i title="English pronunciation respelling">ap-ə-<span style="font-size:90%">TOH</span>-sis</i></a><sup id="cite_ref-ahdictionary_15-0" class="reference"><a href="#cite_note-ahdictionary-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-aboutapop_16-0" class="reference"><a href="#cite_note-aboutapop-16"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>) and the second <i>p</i> pronounced (<span class="rt-commentedText nowrap"><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="/eɪ/: 'a' in 'face'">eɪ</span><span title="'p' in 'pie'">p</span><span title="/ə/: 'a' in 'about'">ə</span><span title="'p' in 'pie'">p</span><span title="/ˈ/: primary stress follows">ˈ</span><span title="'t' in 'tie'">t</span><span title="/oʊ/: 'o' in 'code'">oʊ</span><span title="'s' in 'sigh'">s</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="'s' in 'sigh'">s</span></span>/</a></span></span>).<sup id="cite_ref-ahdictionary_15-1" class="reference"><a href="#cite_note-ahdictionary-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-webster.com_17-0" class="reference"><a href="#cite_note-webster.com-17"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> In English, the <i>p</i> of the Greek <i>-pt-</i> <a href="/wiki/Consonant_cluster" title="Consonant cluster">consonant cluster</a> is typically silent at the beginning of a word (e.g. <a href="/wiki/Pterodactylus" title="Pterodactylus">pterodactyl</a>, <a href="/wiki/Ptolemy" title="Ptolemy">Ptolemy</a>), but articulated when used in combining forms preceded by a vowel, as in <a href="/wiki/Helicopter" title="Helicopter">helicopter</a> or the orders of insects: <a href="/wiki/Fly" title="Fly">diptera</a>, <a href="/wiki/Lepidoptera" title="Lepidoptera">lepidoptera</a>, etc. </p><p>In the original Kerr, Wyllie & Currie paper,<sup id="cite_ref-Kerr1972_11-1" class="reference"><a href="#cite_note-Kerr1972-11"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> there is a footnote regarding the pronunciation: </p> <blockquote><p>We are most grateful to Professor James Cormack of the Department of Greek, University of Aberdeen, for suggesting this term. The word "apoptosis" (<span title="Ancient Greek (to 1453)-language text"><span lang="grc">ἀπόπτωσις</span></span>) is used in Greek to describe the "dropping off" or "falling off" of petals from flowers, or leaves from trees. To show the derivation clearly, we propose that the stress should be on the penultimate syllable, the second half of the word being pronounced like "ptosis" (with the "p" silent), which comes from the same root "to fall", and is already used to describe the drooping of the upper eyelid.</p></blockquote> <div class="mw-heading mw-heading2"><h2 id="Activation_mechanisms">Activation mechanisms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=2" title="Edit section: Activation mechanisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File"><a href="/wiki/File:Apoptosis.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/8/86/Apoptosis.png" decoding="async" width="262" height="511" class="mw-file-element" data-file-width="262" data-file-height="511" /></a><figcaption></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Control_Of_The_Apoptosis_Mecanisms.pdf" class="mw-file-description"><img alt="Control Of The Apoptotic Mechanisms" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Control_Of_The_Apoptosis_Mecanisms.pdf/page1-220px-Control_Of_The_Apoptosis_Mecanisms.pdf.jpg" decoding="async" width="220" height="311" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Control_Of_The_Apoptosis_Mecanisms.pdf/page1-330px-Control_Of_The_Apoptosis_Mecanisms.pdf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/83/Control_Of_The_Apoptosis_Mecanisms.pdf/page1-440px-Control_Of_The_Apoptosis_Mecanisms.pdf.jpg 2x" data-file-width="1239" data-file-height="1754" /></a><figcaption>Control of the apoptotic mechanisms</figcaption></figure> <p>The initiation of apoptosis is tightly regulated by activation mechanisms, because once apoptosis has begun, it inevitably leads to the death of the cell.<sup id="cite_ref-FOOTNOTEAlberts1029_18-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts1029-18"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid14499155_2-1" class="reference"><a href="#cite_note-pmid14499155-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The two best-understood activation mechanisms are the intrinsic pathway (also called the <a href="/wiki/Mitochondrial" class="mw-redirect" title="Mitochondrial">mitochondrial</a> pathway) and the extrinsic pathway.<sup id="cite_ref-FOOTNOTEAlberts1023_19-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts1023-19"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The <i>intrinsic pathway</i> is activated by intracellular signals generated when cells are stressed and depends on the release of proteins from the intermembrane space of mitochondria.<sup id="cite_ref-FOOTNOTEAlberts1032_20-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts1032-20"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> The <i>extrinsic pathway</i> is activated by extracellular ligands binding to cell-surface death receptors, which leads to the formation of the <a href="/wiki/Death-inducing_signaling_complex" title="Death-inducing signaling complex">death-inducing signaling complex</a> (DISC).<sup id="cite_ref-FOOTNOTEAlberts1024_21-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts1024-21"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>A cell initiates intracellular apoptotic signaling in response to a stress,<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> which may bring about cell death. The binding of nuclear receptors by <a href="/wiki/Glucocorticoid" title="Glucocorticoid">glucocorticoids</a>,<sup id="cite_ref-robspath_23-0" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> heat,<sup id="cite_ref-robspath_23-1" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> radiation,<sup id="cite_ref-robspath_23-2" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> nutrient deprivation,<sup id="cite_ref-robspath_23-3" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> viral infection,<sup id="cite_ref-robspath_23-4" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Hypoxia_(medical)" class="mw-redirect" title="Hypoxia (medical)">hypoxia</a>,<sup id="cite_ref-robspath_23-5" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> increased intracellular concentration of free fatty acids<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> and increased intracellular <a href="/wiki/Calcium" title="Calcium">calcium</a> concentration,<sup id="cite_ref-pmid14647298_25-0" class="reference"><a href="#cite_note-pmid14647298-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19898892_26-0" class="reference"><a href="#cite_note-pmid19898892-26"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> for example, by damage to the membrane, can all trigger the release of intracellular apoptotic signals by a damaged cell. A number of cellular components, such as <a href="/wiki/Poly_ADP_ribose_polymerase" class="mw-redirect" title="Poly ADP ribose polymerase">poly ADP ribose polymerase</a>, may also help regulate apoptosis.<sup id="cite_ref-parp1_27-0" class="reference"><a href="#cite_note-parp1-27"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Single cell fluctuations have been observed in experimental studies of stress induced apoptosis.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Before the actual process of cell death is precipitated by enzymes, apoptotic signals must cause regulatory proteins to initiate the apoptosis pathway. This step allows those signals to cause cell death, or the process to be stopped, should the cell no longer need to die. Several proteins are involved, but two main methods of regulation have been identified: the targeting of <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> functionality,<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> or directly transducing the signal via <a href="/wiki/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">adaptor proteins</a> to the apoptotic mechanisms. An extrinsic pathway for initiation identified in several toxin studies is an increase in calcium concentration within a cell caused by drug activity, which also can cause apoptosis via a calcium binding protease <a href="/wiki/Calpain" title="Calpain">calpain</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Intrinsic_pathway">Intrinsic pathway</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=3" title="Edit section: Intrinsic pathway"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The intrinsic pathway is also known as the mitochondrial pathway. <a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondria</a> are essential to multicellular life. Without them, a cell ceases to <a href="/wiki/Cellular_respiration#Aerobic_respiration" title="Cellular respiration">respire aerobically</a> and quickly dies. This fact forms the basis for some apoptotic pathways. Apoptotic proteins that target mitochondria affect them in different ways. They may cause mitochondrial swelling through the formation of membrane pores, or they may increase the permeability of the mitochondrial membrane and cause apoptotic effectors to leak out.<sup id="cite_ref-robspath_23-6" class="reference"><a href="#cite_note-robspath-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> There is also a growing body of evidence indicating that <a href="/wiki/Nitric_oxide" title="Nitric oxide">nitric oxide</a> is able to induce apoptosis by helping to dissipate the <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a> of mitochondria and therefore make it more permeable.<sup id="cite_ref-NO_32-0" class="reference"><a href="#cite_note-NO-32"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Nitric oxide has been implicated in initiating and inhibiting apoptosis through its possible action as a signal molecule of subsequent pathways that activate apoptosis.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>During apoptosis, <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome <i>c</i></a> is released from mitochondria through the actions of the proteins <a href="/wiki/Bcl-2-associated_X_protein" class="mw-redirect" title="Bcl-2-associated X protein">Bax</a> and <a href="/wiki/Bcl-2_homologous_antagonist_killer" title="Bcl-2 homologous antagonist killer">Bak</a>. The mechanism of this release is enigmatic, but appears to stem from a multitude of Bax/Bak homo- and hetero-dimers of Bax/Bak inserted into the outer membrane.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Once cytochrome <i>c</i> is released it binds with Apoptotic protease activating factor – 1 (<i><a href="/wiki/Apaf-1" class="mw-redirect" title="Apaf-1">Apaf-1</a></i>) and <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a>, which then bind to <i>pro-caspase-9</i> to create a protein complex known as an <a href="/wiki/Apoptosome" title="Apoptosome">apoptosome</a>. The apoptosome cleaves the pro-caspase to its active form of <a href="/wiki/Caspase-9" title="Caspase-9">caspase-9</a>, which in turn cleaves and activates pro-caspase into the effector <i>caspase-3</i>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p><p>Mitochondria also release proteins known as SMACs (second mitochondria-derived activator of <a href="/wiki/Caspase" title="Caspase">caspases</a>) into the cell's <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> following the increase in permeability of the mitochondria membranes. SMAC binds to <i><a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">proteins that inhibit apoptosis</a></i> (IAPs) thereby deactivating them, and preventing the IAPs from arresting the process and therefore allowing apoptosis to proceed. IAP also normally suppresses the activity of a group of <a href="/wiki/Cysteine_protease" title="Cysteine protease">cysteine proteases</a> called <a href="/wiki/Caspase" title="Caspase">caspases</a>,<sup id="cite_ref-caspcontrol_35-0" class="reference"><a href="#cite_note-caspcontrol-35"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> which carry out the degradation of the cell. Therefore, the actual degradation enzymes can be seen to be indirectly regulated by mitochondrial permeability.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Extrinsic_pathway">Extrinsic pathway</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=4" title="Edit section: Extrinsic pathway"><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.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/Signal_transduction_pathways.png/500px-Signal_transduction_pathways.png" decoding="async" width="500" height="367" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/Signal_transduction_pathways.png/750px-Signal_transduction_pathways.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fb/Signal_transduction_pathways.png/1000px-Signal_transduction_pathways.png 2x" data-file-width="1858" data-file-height="1364" /></a><figcaption>Overview of signal transduction pathways</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:508px;max-width:508px"><div class="trow"><div class="tsingle" style="width:252px;max-width:252px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:TFN-signalling.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/TFN-signalling.svg/250px-TFN-signalling.svg.png" decoding="async" width="250" height="186" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/TFN-signalling.svg/375px-TFN-signalling.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a8/TFN-signalling.svg/500px-TFN-signalling.svg.png 2x" data-file-width="514" data-file-height="382" /></a></span></div></div><div class="tsingle" style="width:252px;max-width:252px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Fas-signalling.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Fas-signalling.svg/250px-Fas-signalling.svg.png" decoding="async" width="250" height="186" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/Fas-signalling.svg/375px-Fas-signalling.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/60/Fas-signalling.svg/500px-Fas-signalling.svg.png 2x" data-file-width="514" data-file-height="382" /></a></span></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">Overview of TNF (left) and Fas (right) signalling in apoptosis, an example of direct signal transduction</div></div></div></div> <p>Two theories of the direct initiation of apoptotic mechanisms in mammals have been suggested: the <i>TNF-induced</i> (<a href="/wiki/Tumor_necrosis_factor" title="Tumor necrosis factor">tumor necrosis factor</a>) model and the <i>Fas-Fas <a href="/wiki/Ligand" title="Ligand">ligand</a>-mediated</i> model, both involving receptors of the <i>TNF receptor</i> (TNFR) family<sup id="cite_ref-fas_36-0" class="reference"><a href="#cite_note-fas-36"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> coupled to extrinsic signals. </p> <div class="mw-heading mw-heading4"><h4 id="TNF_pathway">TNF pathway</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=5" title="Edit section: TNF pathway"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/TNF-alpha" class="mw-redirect" title="TNF-alpha">TNF-alpha</a> is a <a href="/wiki/Cytokine" title="Cytokine">cytokine</a> produced mainly by activated <a href="/wiki/Macrophage" title="Macrophage">macrophages</a>, and is the major extrinsic mediator of apoptosis. Most cells in the human body have two receptors for TNF-alpha: <a href="/wiki/TNFR1" class="mw-redirect" title="TNFR1">TNFR1</a> and <a href="/wiki/TNFR2" class="mw-redirect" title="TNFR2">TNFR2</a>. The binding of TNF-alpha to TNFR1 has been shown to initiate the pathway that leads to caspase activation via the intermediate membrane proteins TNF receptor-associated death domain (<a href="/wiki/TRADD" title="TRADD">TRADD</a>) and Fas-associated death domain protein (<a href="/wiki/FADD" title="FADD">FADD</a>). <a href="/wiki/CIAP1" class="mw-redirect" title="CIAP1">cIAP1</a>/2 can inhibit TNF-α signaling by binding to <a href="/wiki/TRAF2" title="TRAF2">TRAF2</a>. <a href="/wiki/CFLAR" title="CFLAR">FLIP</a> inhibits the activation of caspase-8.<sup id="cite_ref-tnfr1_37-0" class="reference"><a href="#cite_note-tnfr1-37"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Binding of this receptor can also indirectly lead to the activation of <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> involved in cell survival and inflammatory responses.<sup id="cite_ref-tfnpathway_38-0" class="reference"><a href="#cite_note-tfnpathway-38"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> However, signalling through TNFR1 might also induce apoptosis in a caspase-independent manner.<sup id="cite_ref-LAPFapoptosis_39-0" class="reference"><a href="#cite_note-LAPFapoptosis-39"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template noprint noexcerpt Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:NOTRS" class="mw-redirect" title="Wikipedia:NOTRS"><span title="This claim needs references to better sources. (October 2024)">better source needed</span></a></i>]</sup> The link between TNF-alpha and apoptosis shows why an abnormal production of TNF-alpha plays a fundamental role in several human diseases, especially in <a href="/wiki/Autoimmune_disease" title="Autoimmune disease">autoimmune diseases</a>. The <a href="/wiki/TNF_receptor_superfamily" title="TNF receptor superfamily">TNF-alpha receptor superfamily</a> also includes death receptors (DRs), such as <a href="/wiki/Death_receptor_4" title="Death receptor 4">DR4</a> and <a href="/wiki/Death_receptor_5" title="Death receptor 5">DR5</a>. These receptors bind to the protein <a href="/wiki/TRAIL" title="TRAIL">TRAIL</a> and mediate apoptosis. Apoptosis is known to be one of the primary mechanisms of targeted cancer therapy.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Luminescent iridium complex-peptide hybrids (IPHs) have recently been designed, which mimic TRAIL and bind to death receptors on cancer cells, thereby inducing their apoptosis.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Fas_pathway">Fas pathway</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=6" title="Edit section: Fas pathway"><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/Activation-induced_cell_death" title="Activation-induced cell death">Activation-induced cell death</a></div> <p>The <a href="/wiki/Fas_receptor" title="Fas receptor">fas receptor</a> (First apoptosis signal) – (also known as <i>Apo-1</i> or <i>CD95</i>) is a <a href="/wiki/Transmembrane_protein" title="Transmembrane protein">transmembrane protein</a> of the TNF family which binds the <a href="/wiki/FAS_ligand" class="mw-redirect" title="FAS ligand">Fas ligand</a> (FasL).<sup id="cite_ref-fas_36-1" class="reference"><a href="#cite_note-fas-36"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> The interaction between Fas and FasL results in the formation of the <i>death-inducing signaling complex</i> (DISC), which contains the FADD, caspase-8 and caspase-10. In some types of cells (type I), processed caspase-8 directly activates other members of the caspase family, and triggers the execution of apoptosis of the cell. In other types of cells (type II), the <i>Fas</i>-DISC starts a feedback loop that spirals into increasing release of proapoptotic factors from mitochondria and the amplified activation of caspase-8.<sup id="cite_ref-fassignal_42-0" class="reference"><a href="#cite_note-fassignal-42"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Common_components">Common components</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=7" title="Edit section: Common components"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Following <i>TNF-R1</i> and <i>Fas</i> activation in mammalian cells<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This is only relevant Type II (pancreatic B-cells and hemaetapoetic stem cells), FAS depdnent apoptosis is indepdnent of MOMP (October 2020)">citation needed</span></a></i>]</sup> a balance between proapoptotic (<a href="/wiki/Bcl-2-associated_X_protein" class="mw-redirect" title="Bcl-2-associated X protein">BAX</a>,<sup id="cite_ref-bax_43-0" class="reference"><a href="#cite_note-bax-43"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> <a href="/wiki/BH3_interacting_domain_death_agonist" class="mw-redirect" title="BH3 interacting domain death agonist">BID</a>, <a href="/wiki/Bcl-2_homologous_antagonist_killer" title="Bcl-2 homologous antagonist killer">BAK</a>, or <a href="/wiki/Bcl-2-associated_death_promoter" title="Bcl-2-associated death promoter">BAD</a>) and anti-apoptotic (<i><a href="/wiki/Bcl-Xl" class="mw-redirect" title="Bcl-Xl">Bcl-Xl</a></i> and <i><a href="/wiki/Bcl-2" title="Bcl-2">Bcl-2</a></i>) members of the <i>Bcl-2</i> family are established. This balance is the proportion of proapoptotic <a href="/wiki/Protein_dimer" title="Protein dimer">homodimers</a> that form in the outer-membrane of the mitochondrion. The proapoptotic homodimers are required to make the mitochondrial membrane permeable for the release of caspase activators such as cytochrome c and SMAC. Control of proapoptotic proteins under normal cell conditions of nonapoptotic cells is incompletely understood, but in general, Bax or Bak are activated by the activation of BH3-only proteins, part of the <a href="/wiki/Bcl-2" title="Bcl-2">Bcl-2</a> family.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Caspases">Caspases</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=8" title="Edit section: Caspases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Caspase" title="Caspase">Caspases</a> play the central role in the transduction of ER apoptotic signals. Caspases are proteins that are highly conserved, cysteine-dependent aspartate-specific proteases. There are two types of caspases: initiator caspases (caspases 2, 8, 9, 10, 11, and 12) and effector caspases (caspases 3, 6, and 7). The activation of initiator caspases requires binding to specific oligomeric <a href="/wiki/APAF1" title="APAF1">activator protein</a>. Effector caspases are then activated by these active initiator caspases through <a href="/wiki/Proteolysis" title="Proteolysis">proteolytic</a> cleavage. The active effector caspases then proteolytically degrade a host of intracellular proteins to carry out the cell death program.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading4"><h4 id="Caspase-independent_apoptotic_pathway">Caspase-independent apoptotic pathway</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=9" title="Edit section: Caspase-independent apoptotic pathway"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There also exists a caspase-independent apoptotic pathway that is mediated by AIF (<a href="/wiki/Apoptosis-inducing_factor" title="Apoptosis-inducing factor">apoptosis-inducing factor</a>).<sup id="cite_ref-pmid9989411_45-0" class="reference"><a href="#cite_note-pmid9989411-45"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Apoptosis_model_in_amphibians">Apoptosis model in amphibians</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=10" title="Edit section: Apoptosis model in amphibians"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The frog <i><a href="/wiki/Xenopus_laevis" class="mw-redirect" title="Xenopus laevis">Xenopus laevis</a></i> serves as an ideal model system for the study of the mechanisms of apoptosis. In fact, iodine and thyroxine also stimulate the spectacular apoptosis of the cells of the larval gills, tail and fins in amphibian's metamorphosis, and stimulate the evolution of their nervous system transforming the aquatic, vegetarian tadpole into the terrestrial, carnivorous <a href="/wiki/Frog" title="Frog">frog</a>.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Negative_regulators_of_apoptosis">Negative regulators of apoptosis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=11" title="Edit section: Negative regulators of apoptosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Negative regulation of apoptosis inhibits cell death signaling pathways, helping tumors to evade cell death and developing <a href="/wiki/Drug_resistance" title="Drug resistance">drug resistance</a>. The ratio between anti-apoptotic (Bcl-2) and pro-apoptotic (Bax) proteins determines whether a cell lives or dies.<sup id="cite_ref-pmid31380246_50-0" class="reference"><a href="#cite_note-pmid31380246-50"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid29149100_51-0" class="reference"><a href="#cite_note-pmid29149100-51"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Many families of proteins act as negative regulators categorized into either antiapoptotic factors, such as <a href="/wiki/Inhibitor_of_apoptosis_protein" class="mw-redirect" title="Inhibitor of apoptosis protein">IAPs</a> and <a href="/wiki/Bcl-2_family" title="Bcl-2 family">Bcl-2</a> proteins or prosurvival factors like <a href="/wiki/CFLAR" title="CFLAR">cFLIP</a>, <a href="/wiki/BNIP3" title="BNIP3">BNIP3</a>, <a href="/wiki/FADD" title="FADD">FADD</a>, <a href="/wiki/Protein_kinase_B" title="Protein kinase B">Akt</a>, and <a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-κB</a>.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Proteolytic_caspase_cascade:_Killing_the_cell">Proteolytic caspase cascade: Killing the cell</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=12" title="Edit section: Proteolytic caspase cascade: Killing the cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many pathways and signals lead to apoptosis, but these converge on a single mechanism that actually causes the death of the cell. After a cell receives stimulus, it undergoes organized degradation of cellular organelles by activated proteolytic <a href="/wiki/Caspase" title="Caspase">caspases</a>. In addition to the destruction of cellular organelles, <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> is rapidly and globally degraded by a mechanism that is not yet fully characterized.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> mRNA decay is triggered very early in apoptosis. </p><p>A cell undergoing apoptosis shows a series of characteristic morphological changes. Early alterations include: </p> <ol><li>Cell shrinkage and rounding occur because of the retraction of <a href="/wiki/Lamellipodia" class="mw-redirect" title="Lamellipodia">lamellipodia</a> and the breakdown of the proteinaceous cytoskeleton by caspases.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup></li> <li>The cytoplasm appears dense, and the organelles appear tightly packed.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup></li> <li>Chromatin undergoes condensation into compact patches against the <a href="/wiki/Nuclear_membrane" class="mw-redirect" title="Nuclear membrane">nuclear envelope</a> (also known as the perinuclear envelope) in a process known as <a href="/wiki/Pyknosis" title="Pyknosis">pyknosis</a>, a hallmark of apoptosis.<sup id="cite_ref-nuclearapopt_55-0" class="reference"><a href="#cite_note-nuclearapopt-55"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-chromatindegrad_56-0" class="reference"><a href="#cite_note-chromatindegrad-56"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup></li> <li>The nuclear envelope becomes discontinuous and the DNA inside it is fragmented in a process referred to as <a href="/wiki/Karyorrhexis" title="Karyorrhexis">karyorrhexis</a>. The nucleus breaks into several discrete <i>chromatin bodies</i> or <i>nucleosomal units</i> due to the degradation of DNA.<sup id="cite_ref-nuclearfrag_57-0" class="reference"><a href="#cite_note-nuclearfrag-57"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup></li></ol> <p>Apoptosis progresses quickly and its products are quickly removed, making it difficult to detect or visualize on classical histology sections. During karyorrhexis, <a href="/wiki/Endonuclease" title="Endonuclease">endonuclease</a> activation leaves short DNA fragments, regularly spaced in size. These give a characteristic "laddered" appearance on <a href="/wiki/Agar" title="Agar">agar</a> gel after <a href="/wiki/Electrophoresis" title="Electrophoresis">electrophoresis</a>.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Tests for <a href="/wiki/DNA_laddering" title="DNA laddering">DNA laddering</a> differentiate apoptosis from <a href="/wiki/Ischemia" title="Ischemia">ischemic</a> or toxic cell death.<sup id="cite_ref-Iwata_59-0" class="reference"><a href="#cite_note-Iwata-59"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Apoptotic_cell_disassembly">Apoptotic cell disassembly</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=13" title="Edit section: Apoptotic cell disassembly"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Apoptotic_cell_disassembly.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Apoptotic_cell_disassembly.png/500px-Apoptotic_cell_disassembly.png" decoding="async" width="500" height="216" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Apoptotic_cell_disassembly.png/750px-Apoptotic_cell_disassembly.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/33/Apoptotic_cell_disassembly.png/1000px-Apoptotic_cell_disassembly.png 2x" data-file-width="1333" data-file-height="575" /></a><figcaption>Different steps in apoptotic cell disassembly<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Before the apoptotic cell is disposed of, there is a process of disassembly. There are three recognized steps in apoptotic cell disassembly:<sup id="cite_ref-pmid28102458_61-0" class="reference"><a href="#cite_note-pmid28102458-61"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <ol><li>Membrane blebbing: The cell membrane shows irregular buds known as <a href="/wiki/Bleb_(cell_biology)" title="Bleb (cell biology)">blebs</a>. Initially these are smaller surface blebs. Later these can grow into larger so-called dynamic membrane blebs.<sup id="cite_ref-pmid28102458_61-1" class="reference"><a href="#cite_note-pmid28102458-61"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> An important regulator of apoptotic cell membrane blebbing is <a href="/wiki/ROCK1" title="ROCK1">ROCK1</a> (rho associated coiled-coil-containing protein kinase 1).<sup id="cite_ref-pmid11283606_62-0" class="reference"><a href="#cite_note-pmid11283606-62"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11283607_63-0" class="reference"><a href="#cite_note-pmid11283607-63"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup></li> <li>Formation of membrane protrusions: Some cell types, under specific conditions, may develop different types of long, thin extensions of the cell membrane called membrane protrusions. Three types have been described: <a href="/wiki/Microtubule" title="Microtubule">microtubule</a> spikes, <b>apoptopodia</b> (<i>feet of death</i>), and <b>beaded apoptopodia</b> (the latter having a beads-on-a-string appearance).<sup id="cite_ref-pmid16723742_64-0" class="reference"><a href="#cite_note-pmid16723742-64"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24646995_65-0" class="reference"><a href="#cite_note-pmid24646995-65"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid26074490_66-0" class="reference"><a href="#cite_note-pmid26074490-66"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Pannexin_1" class="mw-redirect" title="Pannexin 1">Pannexin 1</a> is an important component of membrane channels involved in the formation of apoptopodia and beaded apoptopodia.<sup id="cite_ref-pmid24646995_65-1" class="reference"><a href="#cite_note-pmid24646995-65"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Fragmentation_(cell_biology)" title="Fragmentation (cell biology)">Fragmentation</a>: The cell breaks apart into multiple <a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">vesicles</a> called <i>apoptotic bodies</i>, which undergo <a href="/wiki/Phagocytosis" title="Phagocytosis">phagocytosis</a>. The plasma membrane protrusions may help bring apoptotic bodies closer to phagocytes.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup></li></ol> <div class="mw-heading mw-heading3"><h3 id="Removal_of_dead_cells">Removal of dead cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=14" title="Edit section: Removal of dead cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The removal of dead cells by neighboring phagocytic cells has been termed <a href="/wiki/Efferocytosis" title="Efferocytosis">efferocytosis</a>.<sup id="cite_ref-pmid16778289_67-0" class="reference"><a href="#cite_note-pmid16778289-67"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Dying cells that undergo the final stages of apoptosis display phagocytotic molecules, such as <a href="/wiki/Phosphatidylserine" title="Phosphatidylserine">phosphatidylserine</a>, on their cell surface.<sup id="cite_ref-Phosphatidylserine_68-0" class="reference"><a href="#cite_note-Phosphatidylserine-68"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> Phosphatidylserine is normally found on the inner leaflet surface of the plasma membrane, but is redistributed during apoptosis to the extracellular surface by a protein known as <a href="/wiki/Phospholipid_scramblase" title="Phospholipid scramblase">scramblase</a>.<sup id="cite_ref-phago2_69-0" class="reference"><a href="#cite_note-phago2-69"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> These molecules mark the cell for <a href="/wiki/Phagocytosis" title="Phagocytosis">phagocytosis</a> by cells possessing the appropriate receptors, such as macrophages.<sup id="cite_ref-phago1_70-0" class="reference"><a href="#cite_note-phago1-70"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> The removal of dying cells by phagocytes occurs in an orderly manner without eliciting an <a href="/wiki/Inflammatory_response" class="mw-redirect" title="Inflammatory response">inflammatory response</a>.<sup id="cite_ref-phago1B_71-0" class="reference"><a href="#cite_note-phago1B-71"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> During apoptosis cellular RNA and DNA are separated from each other and sorted to different apoptotic bodies; separation of RNA is initiated as nucleolar segregation.<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Pathway_knock-outs">Pathway knock-outs</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=15" title="Edit section: Pathway knock-outs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many <a href="/wiki/Gene_knockout" title="Gene knockout">knock-outs</a> have been made in the apoptosis pathways to test the function of each of the proteins. Several caspases, in addition to <a href="/wiki/APAF1" title="APAF1">APAF1</a> and <a href="/wiki/FADD" title="FADD">FADD</a>, have been mutated to determine the new phenotype. In order to create a tumor necrosis factor (TNF) knockout, an exon containing the nucleotides 3704–5364 was removed from the gene.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> This exon encodes a portion of the mature TNF domain, as well as the leader sequence, which is a highly conserved region necessary for proper intracellular processing. TNF-/- mice develop normally and have no gross structural or morphological abnormalities. However, upon immunization with SRBC (sheep red blood cells), these mice demonstrated a deficiency in the maturation of an antibody response; they were able to generate normal levels of IgM, but could not develop specific IgG levels.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Apaf-1 is the protein that turns on caspase 9 by cleavage to begin the caspase cascade that leads to apoptosis.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> Since a -/- mutation in the APAF-1 gene is embryonic lethal, a gene trap strategy was used in order to generate an APAF-1 -/- mouse. This assay is used to disrupt gene function by creating an intragenic gene fusion. When an APAF-1 gene trap is introduced into cells, many morphological changes occur, such as spina bifida, the persistence of interdigital webs, and open brain.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> In addition, after embryonic day 12.5, the brain of the embryos showed several structural changes. APAF-1 cells are protected from apoptosis stimuli such as irradiation. A BAX-1 knock-out mouse exhibits normal forebrain formation and a decreased programmed cell death in some neuronal populations and in the spinal cord, leading to an increase in motor neurons.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p>The caspase proteins are integral parts of the apoptosis pathway, so it follows that knock-outs made have varying damaging results. A caspase 9 knock-out leads to a severe brain malformation <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (October 2022)">citation needed</span></a></i>]</sup>. A caspase 8 knock-out leads to cardiac failure and thus embryonic lethality <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (October 2022)">citation needed</span></a></i>]</sup>. However, with the use of cre-lox technology, a caspase 8 knock-out has been created that exhibits an increase in peripheral T cells, an impaired T cell response, and a defect in neural tube closure <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (October 2022)">citation needed</span></a></i>]</sup>. These mice were found to be resistant to apoptosis mediated by CD95, TNFR, etc. but not resistant to apoptosis caused by UV irradiation, chemotherapeutic drugs, and other stimuli. Finally, a caspase 3 knock-out was characterized by ectopic cell masses in the brain and abnormal apoptotic features such as membrane blebbing or nuclear fragmentation <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (October 2022)">citation needed</span></a></i>]</sup>. A remarkable feature of these KO mice is that they have a very restricted phenotype: Casp3, 9, APAF-1 KO mice have deformations of neural tissue and FADD and Casp 8 KO showed defective heart development, however, in both types of KO other organs developed normally and some cell types were still sensitive to apoptotic stimuli suggesting that unknown proapoptotic pathways exist.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Methods_for_distinguishing_apoptotic_from_necrotic_cells">Methods for distinguishing apoptotic from necrotic cells</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=16" title="Edit section: Methods for distinguishing apoptotic from necrotic cells"><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:Apoptosis_in_mouse_pre-adipocytes.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Apoptosis_in_mouse_pre-adipocytes.gif/220px-Apoptosis_in_mouse_pre-adipocytes.gif" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/a/a0/Apoptosis_in_mouse_pre-adipocytes.gif 1.5x" data-file-width="300" data-file-height="300" /></a><figcaption>Long-term live cell imaging (12h) of multinucleated mouse pre-Adipocyte trying to undergo mitosis. Due to the excess of genetic material the cell fails to replicate and dies by apoptosis.</figcaption></figure> <p>Label-free <a href="/wiki/Live_cell_imaging" class="mw-redirect" title="Live cell imaging">live cell imaging</a>, <a href="/wiki/Time-lapse_microscopy" title="Time-lapse microscopy">time-lapse microscopy</a>, <a href="/wiki/Flow_cytometry" title="Flow cytometry">flow fluorocytometry</a>, and <a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">transmission electron microscopy</a> can be used to compare apoptotic and necrotic cells. There are also various biochemical techniques for analysis of cell surface markers (phosphatidylserine exposure versus cell permeability by flow cytometry), cellular markers such as <a href="/wiki/DNA_fragmentation" title="DNA fragmentation">DNA fragmentation</a><sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> (flow cytometry),<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> caspase activation, Bid cleavage, and cytochrome c release (<a href="/wiki/Western_blot" title="Western blot">Western blotting</a>). Supernatant screening for caspases, HMGB1, and cytokeratin 18 release can identify primary from secondary necrotic cells. However, no distinct surface or biochemical markers of necrotic cell death have been identified yet, and only negative markers are available. These include absence of apoptotic markers (caspase activation, cytochrome c release, and oligonucleosomal DNA fragmentation) and differential kinetics of cell death markers (phosphatidylserine exposure and cell membrane permeabilization). A selection of techniques that can be used to distinguish apoptosis from necroptotic cells could be found in these references.<sup id="cite_ref-methods_80-0" class="reference"><a href="#cite_note-methods-80"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-methods2_81-0" class="reference"><a href="#cite_note-methods2-81"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-methods3_82-0" class="reference"><a href="#cite_note-methods3-82"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Implication_in_disease">Implication in disease</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=17" title="Edit section: Implication in disease"><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:Apoptosis_multi_mouseliver.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Apoptosis_multi_mouseliver.jpg/220px-Apoptosis_multi_mouseliver.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Apoptosis_multi_mouseliver.jpg/330px-Apoptosis_multi_mouseliver.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/d/d3/Apoptosis_multi_mouseliver.jpg 2x" data-file-width="417" data-file-height="417" /></a><figcaption>A section of mouse liver showing several apoptotic cells, indicated by arrows</figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Apoptosis_stained.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Apoptosis_stained.jpg/220px-Apoptosis_stained.jpg" decoding="async" width="220" height="216" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Apoptosis_stained.jpg/330px-Apoptosis_stained.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Apoptosis_stained.jpg/440px-Apoptosis_stained.jpg 2x" data-file-width="485" data-file-height="477" /></a><figcaption>A section of mouse liver <a href="/wiki/Staining_(biology)" class="mw-redirect" title="Staining (biology)">stained</a> to show cells undergoing apoptosis (orange)</figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Apoptosis.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Apoptosis.jpg/220px-Apoptosis.jpg" decoding="async" width="220" height="161" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Apoptosis.jpg/330px-Apoptosis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/08/Apoptosis.jpg/440px-Apoptosis.jpg 2x" data-file-width="2657" data-file-height="1944" /></a><figcaption>Neonatal cardiomyocytes ultrastructure after anoxia-reoxygenation</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Defective_pathways">Defective pathways</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=18" title="Edit section: Defective pathways"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The many different types of apoptotic pathways contain a multitude of different biochemical components, many of them not yet understood.<sup id="cite_ref-pathogenesis_84-0" class="reference"><a href="#cite_note-pathogenesis-84"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> As a pathway is more or less sequential in nature, removing or modifying one component leads to an effect in another. In a living organism, this can have disastrous effects, often in the form of disease or disorder. A discussion of every disease caused by modification of the various apoptotic pathways would be impractical, but the concept overlying each one is the same: The normal functioning of the pathway has been disrupted in such a way as to impair the ability of the cell to undergo normal apoptosis. This results in a cell that lives past its "use-by date" and is able to replicate and pass on any faulty machinery to its progeny, increasing the likelihood of the cell's becoming cancerous or diseased.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p><p>A recently described example of this concept in action can be seen in the development of a lung cancer called <a href="/wiki/NCI-H460" class="mw-redirect" title="NCI-H460">NCI-H460</a>.<sup id="cite_ref-h460_85-0" class="reference"><a href="#cite_note-h460-85"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> The <i>X-linked inhibitor of apoptosis protein</i> (<a href="/wiki/XIAP" title="XIAP">XIAP</a>) is <a href="/wiki/Gene_expression" title="Gene expression">overexpressed</a> in cells of the H460 <a href="/wiki/Cell_line" class="mw-redirect" title="Cell line">cell line</a>. XIAPs bind to the processed form of caspase-9 and suppress the activity of apoptotic activator <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome c</a>, therefore overexpression leads to a decrease in the number of proapoptotic agonists. As a consequence, the balance of anti-apoptotic and proapoptotic effectors is upset in favour of the former, and the damaged cells continue to replicate despite being directed to die. Defects in regulation of apoptosis in cancer cells occur often at the level of control of transcription factors. As a particular example, defects in molecules that control transcription factor NF-κB in cancer change the mode of transcriptional regulation and the response to apoptotic signals, to curtail dependence on the tissue that the cell belongs. This degree of independence from external survival signals, can enable cancer metastasis.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Dysregulation_of_p53">Dysregulation of p53</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=19" title="Edit section: Dysregulation of p53"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The tumor-suppressor protein <a href="/wiki/P53" title="P53">p53</a> accumulates when DNA is damaged due to a chain of biochemical factors. Part of this pathway includes alpha-<a href="/wiki/Interferon" title="Interferon">interferon</a> and beta-interferon, which induce transcription of the <i>p53</i> gene, resulting in the increase of p53 protein level and enhancement of cancer cell-apoptosis.<sup id="cite_ref-takaoka_87-0" class="reference"><a href="#cite_note-takaoka-87"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> p53 prevents the cell from replicating by stopping the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> at G1, or interphase, to give the cell time to repair; however, it will induce apoptosis if damage is extensive and repair efforts fail.<sup id="cite_ref-pmid12052432_88-0" class="reference"><a href="#cite_note-pmid12052432-88"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> Any disruption to the regulation of the <i>p53</i> or interferon genes will result in impaired apoptosis and the possible formation of tumors.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inhibition">Inhibition</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=20" title="Edit section: Inhibition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Inhibition of apoptosis can result in a number of cancers, inflammatory diseases, and viral infections. It was originally believed that the associated accumulation of cells was due to an increase in cellular proliferation, but it is now known that it is also due to a decrease in cell death. The most common of these diseases is cancer, the disease of excessive cellular proliferation, which is often characterized by an overexpression of <a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">IAP</a> family members. As a result, the malignant cells experience an abnormal response to apoptosis induction: Cycle-regulating genes (such as p53, ras or c-myc) are mutated or inactivated in diseased cells, and further genes (such as bcl-2) also modify their expression in tumors. Some apoptotic factors are vital during mitochondrial respiration e.g. cytochrome C.<sup id="cite_ref-iopscience.iop.org_89-0" class="reference"><a href="#cite_note-iopscience.iop.org-89"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> Pathological inactivation of apoptosis in cancer cells is correlated with frequent respiratory metabolic shifts toward glycolysis (an observation known as the "Warburg hypothesis".<sup id="cite_ref-Warburg_1956_90-0" class="reference"><a href="#cite_note-Warburg_1956-90"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="HeLa_cell">HeLa cell</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=21" title="Edit section: HeLa cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Apoptosis in <a href="/wiki/HeLa" title="HeLa">HeLa</a><sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> cells is inhibited by proteins produced by the cell; these inhibitory proteins target retinoblastoma tumor-suppressing proteins.<sup id="cite_ref-DelPuerto_92-0" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> These tumor-suppressing proteins regulate the cell cycle, but are rendered inactive when bound to an inhibitory protein.<sup id="cite_ref-DelPuerto_92-1" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> HPV E6 and E7 are inhibitory proteins expressed by the human papillomavirus, HPV being responsible for the formation of the cervical tumor from which HeLa cells are derived.<sup id="cite_ref-Liu_93-0" class="reference"><a href="#cite_note-Liu-93"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> HPV E6 causes p53, which regulates the cell cycle, to become inactive.<sup id="cite_ref-Niu_94-0" class="reference"><a href="#cite_note-Niu-94"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> HPV E7 binds to retinoblastoma tumor suppressing proteins and limits its ability to control cell division.<sup id="cite_ref-Niu_94-1" class="reference"><a href="#cite_note-Niu-94"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> These two inhibitory proteins are partially responsible for HeLa cells' immortality by inhibiting apoptosis to occur.<sup id="cite_ref-Liu_McKalip_95-0" class="reference"><a href="#cite_note-Liu_McKalip-95"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Treatments">Treatments</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=22" title="Edit section: Treatments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information on a clinical pathology test that measures apoptosis: <a href="/wiki/MiCK_assay" title="MiCK assay">MiCK assay</a></div> <p>The main method of treatment for potential death from signaling-related diseases involves either increasing or decreasing the susceptibility of apoptosis in diseased cells, depending on whether the disease is caused by either the inhibition of or excess apoptosis. For instance, treatments aim to restore apoptosis to treat diseases with deficient cell death and to increase the apoptotic threshold to treat diseases involved with excessive cell death. To stimulate apoptosis, one can increase the number of death receptor ligands (such as TNF or TRAIL), antagonize the anti-apoptotic Bcl-2 pathway, or introduce Smac mimetics to inhibit the inhibitor (IAPs).<sup id="cite_ref-pmid31380246_50-1" class="reference"><a href="#cite_note-pmid31380246-50"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> The addition of agents such as Herceptin, Iressa, or Gleevec works to stop cells from cycling and causes apoptosis activation by blocking growth and survival signaling further upstream. Finally, adding p53-<a href="/wiki/MDM2" class="mw-redirect" title="MDM2">MDM2</a> complexes displaces p53 and activates the p53 pathway, leading to cell cycle arrest and apoptosis. Many different methods can be used either to stimulate or to inhibit apoptosis in various places along the death signaling pathway.<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> </p><p>Apoptosis is a multi-step, multi-pathway cell-death programme that is inherent in every cell of the body. In cancer, the apoptosis cell-division ratio is altered. Cancer treatment by chemotherapy and irradiation kills target cells primarily by inducing apoptosis.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Hyperactive_apoptosis">Hyperactive apoptosis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=23" title="Edit section: Hyperactive apoptosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>On the other hand, loss of control of cell death (resulting in excess apoptosis) can lead to neurodegenerative diseases, hematologic diseases, and tissue damage. Neurons that rely on mitochondrial respiration undergo apoptosis in neurodegenerative diseases such as Alzheimer's<sup id="cite_ref-pmid7704018_97-0" class="reference"><a href="#cite_note-pmid7704018-97"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> and Parkinson's.<sup id="cite_ref-pmid8782165_98-0" class="reference"><a href="#cite_note-pmid8782165-98"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> (an observation known as the "Inverse Warburg hypothesis"<sup id="cite_ref-iopscience.iop.org_89-1" class="reference"><a href="#cite_note-iopscience.iop.org-89"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid25642192_99-0" class="reference"><a href="#cite_note-pmid25642192-99"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup>). Moreover, there is an inverse epidemiological comorbidity between neurodegenerative diseases and cancer.<sup id="cite_ref-pmid23843468_100-0" class="reference"><a href="#cite_note-pmid23843468-100"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> The progression of HIV is directly linked to excess, unregulated apoptosis. In a healthy individual, the number of CD4+ lymphocytes is in balance with the cells generated by the bone marrow; however, in HIV-positive patients, this balance is lost due to an inability of the bone marrow to regenerate CD4+ cells. In the case of HIV, CD4+ lymphocytes die at an accelerated rate through uncontrolled apoptosis, when stimulated. At the molecular level, hyperactive apoptosis can be caused by defects in signaling pathways that regulate the Bcl-2 family proteins. Increased expression of apoptotic proteins such as BIM, or their decreased proteolysis, leads to cell death and can cause a number of pathologies, depending on the cells where excessive activity of BIM occurs. Cancer cells can escape apoptosis through mechanisms that suppress BIM expression or by increased proteolysis of BIM.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="removed citation to predatory publisher content (December 2019)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading4"><h4 id="Treatments_2">Treatments</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=24" title="Edit section: Treatments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Treatments aiming to inhibit works to block specific caspases. Finally, the Akt protein kinase promotes cell survival through two pathways. Akt phosphorylates and inhibits Bad (a Bcl-2 family member), causing Bad to interact with the <a href="/wiki/14-3-3" class="mw-redirect" title="14-3-3">14-3-3</a> scaffold, resulting in Bcl dissociation and thus cell survival. Akt also activates IKKα, which leads to NF-κB activation and cell survival. Active NF-κB induces the expression of anti-apoptotic genes such as Bcl-2, resulting in inhibition of apoptosis. NF-κB has been found to play both an antiapoptotic role and a proapoptotic role depending on the stimuli utilized and the cell type.<sup id="cite_ref-Farhana_2005_101-0" class="reference"><a href="#cite_note-Farhana_2005-101"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="HIV_progression">HIV progression</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=25" title="Edit section: HIV progression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The progression of the <a href="/wiki/Human_immunodeficiency_virus" class="mw-redirect" title="Human immunodeficiency virus">human immunodeficiency virus</a> infection into <a href="/wiki/AIDS" class="mw-redirect" title="AIDS">AIDS</a> is due primarily to the depletion of <a href="/wiki/T_helper_cell" title="T helper cell">CD4+ T-helper lymphocytes</a> in a manner that is too rapid for the body's bone marrow to replenish the cells, leading to a compromised immune system. One of the mechanisms by which T-helper cells are depleted is apoptosis, which results from a series of biochemical pathways:<sup id="cite_ref-Judie_102-0" class="reference"><a href="#cite_note-Judie-102"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> </p> <ol><li>HIV enzymes deactivate anti-apoptotic <i>Bcl-2</i>. This does not directly cause cell death but primes the cell for apoptosis should the appropriate signal be received. In parallel, these enzymes activate proapoptotic <i>procaspase-8</i>, which does directly activate the mitochondrial events of apoptosis.</li> <li>HIV may increase the level of cellular proteins that prompt Fas-mediated apoptosis.</li> <li>HIV proteins decrease the amount of <a href="/wiki/CD4" title="CD4">CD4</a> glycoprotein marker present on the cell membrane.</li> <li>Released viral particles and proteins present in extracellular fluid are able to induce apoptosis in nearby "bystander" T helper cells.</li> <li>HIV decreases the production of molecules involved in marking the cell for apoptosis, giving the virus time to replicate and continue releasing apoptotic agents and virions into the surrounding tissue.</li> <li>The infected CD4+ cell may also receive the death signal from a cytotoxic T cell.</li></ol> <p>Cells may also die as direct consequences of viral infections. HIV-1 expression induces tubular cell G2/M arrest and apoptosis.<sup id="cite_ref-103" class="reference"><a href="#cite_note-103"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> The progression from HIV to AIDS is not immediate or even necessarily rapid; HIV's cytotoxic activity toward CD4+ lymphocytes is classified as AIDS once a given patient's CD4+ cell count falls below 200.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p><p>Researchers from Kumamoto University in Japan have developed a new method to eradicate HIV in viral reservoir cells, named "Lock-in and apoptosis." Using the synthesized compound Heptanoylphosphatidyl L-Inositol Pentakisphophate (or L-Hippo) to bind strongly to the HIV protein PR55Gag, they were able to suppress viral budding. By suppressing viral budding, the researchers were able to trap the HIV virus in the cell and allow for the cell to undergo apoptosis (natural cell death). Associate Professor Mikako Fujita has stated that the approach is not yet available to HIV patients because the research team has to conduct further research on combining the drug therapy that currently exists with this "Lock-in and apoptosis" approach to lead to complete recovery from HIV.<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Viral_infection">Viral infection</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=26" title="Edit section: Viral infection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Viral induction of apoptosis occurs when one or several cells of a living organism are infected with a <a href="/wiki/Virus" title="Virus">virus</a>, leading to cell death. Cell death in organisms is necessary for the normal development of cells and the cell cycle maturation.<sup id="cite_ref-Indran_106-0" class="reference"><a href="#cite_note-Indran-106"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> It is also important in maintaining the regular functions and activities of cells. </p><p>Viruses can trigger apoptosis of infected cells via a range of mechanisms including: </p> <ul><li>Receptor binding</li> <li>Activation of <a href="/wiki/Protein_kinase_R" title="Protein kinase R">protein kinase R</a> (PKR)</li> <li>Interaction with p53</li> <li>Expression of viral proteins coupled to MHC proteins on the surface of the infected cell, allowing recognition by cells of the immune system (such as <a href="/wiki/Natural_killer_cell" title="Natural killer cell">natural killer</a> and <a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">cytotoxic T cells</a>) that then induce the infected cell to undergo apoptosis.<sup id="cite_ref-Everett_107-0" class="reference"><a href="#cite_note-Everett-107"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup></li></ul> <p><a href="/wiki/Canine_distemper_virus" class="mw-redirect" title="Canine distemper virus">Canine distemper virus</a> (CDV) is known to cause apoptosis in central nervous system and lymphoid tissue of infected dogs in vivo and in vitro.<sup id="cite_ref-Nishi_108-0" class="reference"><a href="#cite_note-Nishi-108"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> Apoptosis caused by CDV is typically induced via the <a href="/wiki/Death-inducing_signaling_complex" title="Death-inducing signaling complex">extrinsic pathway</a>, which activates <a href="/wiki/Caspases" class="mw-redirect" title="Caspases">caspases</a> that disrupt cellular function and eventually leads to the cells death.<sup id="cite_ref-DelPuerto_92-2" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> In normal cells, CDV activates caspase-8 first, which works as the initiator protein followed by the executioner protein caspase-3.<sup id="cite_ref-DelPuerto_92-3" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> However, apoptosis induced by CDV in HeLa cells does not involve the initiator protein caspase-8. HeLa cell apoptosis caused by CDV follows a different mechanism than that in vero cell lines.<sup id="cite_ref-DelPuerto_92-4" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> This change in the caspase cascade suggests CDV induces apoptosis via the <a href="/wiki/Intrinsic_apoptosis" class="mw-redirect" title="Intrinsic apoptosis">intrinsic pathway</a>, excluding the need for the initiator caspase-8. The executioner protein is instead activated by the internal stimuli caused by viral infection not a caspase cascade.<sup id="cite_ref-DelPuerto_92-5" class="reference"><a href="#cite_note-DelPuerto-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> </p><p>The <a href="/wiki/Oropouche_virus" class="mw-redirect" title="Oropouche virus">Oropouche virus</a> (OROV) is found in the family <i><a href="/wiki/Bunyaviridae" class="mw-redirect" title="Bunyaviridae">Bunyaviridae</a></i>. The study of apoptosis brought on by <i>Bunyaviridae</i> was initiated in 1996, when it was observed that apoptosis was induced by the <a href="/wiki/La_Crosse_encephalitis" title="La Crosse encephalitis">La Crosse virus</a> into the kidney cells of baby hamsters and into the brains of baby mice.<sup id="cite_ref-Acrani_109-0" class="reference"><a href="#cite_note-Acrani-109"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> </p><p>OROV is a disease that is transmitted between humans by the biting midge (<i><a href="/wiki/Culicoides_paraensis" title="Culicoides paraensis">Culicoides paraensis</a></i>).<sup id="cite_ref-Azevedo_110-0" class="reference"><a href="#cite_note-Azevedo-110"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> It is referred to as a <a href="/wiki/Zoonotic" class="mw-redirect" title="Zoonotic">zoonotic</a> <a href="/wiki/Arbovirus" title="Arbovirus">arbovirus</a> and causes febrile illness, characterized by the onset of a sudden fever known as Oropouche fever.<sup id="cite_ref-Santos_111-0" class="reference"><a href="#cite_note-Santos-111"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> </p><p>The Oropouche virus also causes disruption in cultured cells – cells that are cultivated in distinct and specific conditions. An example of this can be seen in <a href="/wiki/HeLa_cells" class="mw-redirect" title="HeLa cells">HeLa cells</a>, whereby the cells begin to degenerate shortly after they are infected.<sup id="cite_ref-Acrani_109-1" class="reference"><a href="#cite_note-Acrani-109"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> </p><p>With the use of <a href="/wiki/Gel_electrophoresis" title="Gel electrophoresis">gel electrophoresis</a>, it can be observed that OROV causes <a href="/wiki/DNA" title="DNA">DNA</a> fragmentation in HeLa cells. It can be interpreted by counting, measuring, and analyzing the cells of the Sub/G1 cell population.<sup id="cite_ref-Acrani_109-2" class="reference"><a href="#cite_note-Acrani-109"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> When HeLA cells are infected with OROV, the <a href="/wiki/Cytochrome_C" class="mw-redirect" title="Cytochrome C">cytochrome C</a> is released from the membrane of the mitochondria, into the cytosol of the cells. This type of interaction shows that apoptosis is activated via an intrinsic pathway.<sup id="cite_ref-Indran_106-1" class="reference"><a href="#cite_note-Indran-106"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> </p><p>In order for apoptosis to occur within OROV, viral uncoating, viral internalization, along with the replication of cells is necessary. Apoptosis in some viruses is activated by extracellular stimuli. However, studies have demonstrated that the OROV infection causes apoptosis to be activated through intracellular stimuli and involves the mitochondria.<sup id="cite_ref-Acrani_109-3" class="reference"><a href="#cite_note-Acrani-109"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> </p><p>Many viruses encode proteins that can inhibit apoptosis.<sup id="cite_ref-Teodora_112-0" class="reference"><a href="#cite_note-Teodora-112"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> Several viruses encode viral homologs of Bcl-2. These homologs can inhibit proapoptotic proteins such as BAX and BAK, which are essential for the activation of apoptosis. Examples of viral Bcl-2 proteins include the <a href="/wiki/Epstein-Barr_virus" class="mw-redirect" title="Epstein-Barr virus">Epstein-Barr virus</a> BHRF1 protein and the <a href="/wiki/Adenovirus" class="mw-redirect" title="Adenovirus">adenovirus</a> E1B 19K protein.<sup id="cite_ref-Polster_113-0" class="reference"><a href="#cite_note-Polster-113"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> Some viruses express caspase inhibitors that inhibit caspase activity and an example is the CrmA protein of cowpox viruses. Whilst a number of viruses can block the effects of TNF and Fas. For example, the M-T2 protein of myxoma viruses can bind TNF preventing it from binding the TNF receptor and inducing a response.<sup id="cite_ref-Hay_114-0" class="reference"><a href="#cite_note-Hay-114"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> Furthermore, many viruses express p53 inhibitors that can bind p53 and inhibit its transcriptional transactivation activity. As a consequence, p53 cannot induce apoptosis, since it cannot induce the expression of proapoptotic proteins. The adenovirus E1B-55K protein and the <a href="/wiki/Hepatitis_B_virus" title="Hepatitis B virus">hepatitis B virus</a> HBx protein are examples of viral proteins that can perform such a function.<sup id="cite_ref-Wang_115-0" class="reference"><a href="#cite_note-Wang-115"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup> </p><p>Viruses can remain intact from apoptosis in particular in the latter stages of infection. They can be exported in the <i>apoptotic bodies</i> that pinch off from the surface of the dying cell, and the fact that they are engulfed by phagocytes prevents the initiation of a host response. This favours the spread of the virus.<sup id="cite_ref-Hay_114-1" class="reference"><a href="#cite_note-Hay-114"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Prion" title="Prion">Prions</a> can cause apoptosis in <a href="/wiki/Neuron" title="Neuron">neurons</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Plants">Plants</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=27" title="Edit section: Plants"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Programmed_cell_death" title="Programmed cell death">Programmed cell death</a> in plants has a number of molecular similarities to that of animal apoptosis, but it also has differences, notable ones being the presence of a <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a> and the lack of an <a href="/wiki/Immune_system" title="Immune system">immune system</a> that removes the pieces of the dead cell. Instead of an immune response, the dying cell synthesizes substances to break itself down and places them in a <a href="/wiki/Vacuole" title="Vacuole">vacuole</a> that ruptures as the cell dies. Additionally, plants do not contain phagocytic cells, which are essential in the process of breaking down and removing apoptotic bodies.<sup id="cite_ref-116" class="reference"><a href="#cite_note-116"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup> Whether this whole process resembles animal apoptosis closely enough to warrant using the name <i>apoptosis</i> (as opposed to the more general <i>programmed cell death</i>) is unclear.<sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-dickman2017_118-0" class="reference"><a href="#cite_note-dickman2017-118"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Caspase-independent_apoptosis">Caspase-independent apoptosis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=28" title="Edit section: Caspase-independent apoptosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The characterization of the caspases allowed the development of caspase inhibitors, which can be used to determine whether a cellular process involves active caspases. Using these inhibitors it was discovered that cells can die while displaying a morphology similar to apoptosis without caspase activation.<sup id="cite_ref-pmid8962091_119-0" class="reference"><a href="#cite_note-pmid8962091-119"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> Later studies linked this phenomenon to the release of AIF (<a href="/wiki/Apoptosis-inducing_factor" title="Apoptosis-inducing factor">apoptosis-inducing factor</a>) from the mitochondria and its translocation into the nucleus mediated by its NLS (<a href="/wiki/Nuclear_localization_sequence" title="Nuclear localization sequence">nuclear localization signal</a>). Inside the mitochondria, AIF is anchored to the inner membrane. In order to be released, the protein is cleaved by a calcium-dependent <a href="/wiki/Calpain" title="Calpain">calpain protease</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2024)">citation needed</span></a></i>]</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=Apoptosis&action=edit&section=29" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1259569809">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{clear:left;float:left;margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><span class="portalbox-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Issoria_lathonia.jpg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/32px-Issoria_lathonia.jpg" decoding="async" width="32" height="23" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/48px-Issoria_lathonia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/64px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a></span></span><span class="portalbox-link"><a href="/wiki/Portal:Biology" title="Portal:Biology">Biology portal</a></span></li></ul> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 22em;"> <ul><li><a href="/wiki/Anoikis" title="Anoikis">Anoikis</a></li> <li><a href="/wiki/Apaf-1" class="mw-redirect" title="Apaf-1">Apaf-1</a></li> <li><a href="/wiki/Apo2.7" title="Apo2.7">Apo2.7</a></li> <li><a href="/wiki/Apoptotic_DNA_fragmentation" title="Apoptotic DNA fragmentation">Apoptotic DNA fragmentation</a></li> <li><a href="/wiki/Atromentin" title="Atromentin">Atromentin</a> induces apoptosis in human <a href="/wiki/Leukemia" title="Leukemia">leukemia</a> <a href="/wiki/U937_cells" class="mw-redirect" title="U937 cells">U937 cells</a>.<sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Autolysis_(biology)" title="Autolysis (biology)">Autolysis</a></li> <li><a href="/wiki/Autophagy" title="Autophagy">Autophagy</a></li> <li><a href="/wiki/Cisplatin" title="Cisplatin">Cisplatin</a></li> <li><a href="/wiki/Cytotoxicity" title="Cytotoxicity">Cytotoxicity</a></li> <li><a href="/wiki/Entosis" title="Entosis">Entosis</a></li> <li><a href="/wiki/Ferroptosis" title="Ferroptosis">Ferroptosis</a></li> <li><a href="/wiki/Homeostasis" title="Homeostasis">Homeostasis</a></li> <li><a href="/wiki/Immunology" title="Immunology">Immunology</a></li> <li><a href="/wiki/Necrobiosis" title="Necrobiosis">Necrobiosis</a></li> <li><a href="/wiki/Necrosis" title="Necrosis">Necrosis</a></li> <li><a href="/wiki/Necrotaxis" title="Necrotaxis">Necrotaxis</a></li> <li><a href="/wiki/Nemosis" title="Nemosis">Nemosis</a></li> <li><a href="/wiki/Mitotic_catastrophe" title="Mitotic catastrophe">Mitotic catastrophe</a></li> <li><a href="/wiki/P53" title="P53">p53</a></li> <li><a href="/wiki/Paraptosis" title="Paraptosis">Paraptosis</a></li> <li><a href="/wiki/Pseudoapoptosis" title="Pseudoapoptosis">Pseudoapoptosis</a></li> <li><a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="Explanatory_footnotes">Explanatory footnotes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=30" title="Edit section: Explanatory footnotes"><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-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Note that the average human adult has more than 13 trillion cells (<span class="nowrap"><span data-sort-value="7013130000000000000♠"></span>1.3<span style="margin-left:0.25em;margin-right:0.15em;">×</span>10<sup>13</sup></span>),<sup id="cite_ref-FOOTNOTEAlberts2_3-0" class="reference"><a href="#cite_note-FOOTNOTEAlberts2-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> of which at most only 70 billion (<span class="nowrap"><span data-sort-value="7010700000000000000♠"></span>7.0<span style="margin-left:0.25em;margin-right:0.15em;">×</span>10<sup>10</sup></span>) die per day. That is, about 5 out of every 1,000 cells (0.5%) die each day due to apoptosis.</span> </li> <li id="cite_note-91"><span class="mw-cite-backlink"><b><a href="#cite_ref-91">^</a></b></span> <span class="reference-text">HeLa cells are an immortalized cancer cell line used frequently in research. The cell line was established by removing cells directly from <a href="/wiki/Henrietta_Lacks" title="Henrietta Lacks">Henrietta Lacks</a>, a cancer patient.</span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Citations">Citations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=31" title="Edit section: Citations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist reflist-columns references-column-width" style="column-width: 35em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFGreen2011" class="citation book cs1">Green D (2011). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=s8jBcQAACAAJ"><i>Means to an End: Apoptosis and other Cell Death Mechanisms</i></a>. Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-888-1" title="Special:BookSources/978-0-87969-888-1"><bdi>978-0-87969-888-1</bdi></a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200726071718/https://books.google.com/books?id=s8jBcQAACAAJ">Archived</a> from the original on 2020-07-26<span class="reference-accessdate">. Retrieved <span class="nowrap">2020-05-25</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Means+to+an+End%3A+Apoptosis+and+other+Cell+Death+Mechanisms&rft.place=Cold+Spring+Harbor%2C+NY&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=2011&rft.isbn=978-0-87969-888-1&rft.aulast=Green&rft.aufirst=D&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Ds8jBcQAACAAJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-pmid14499155-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid14499155_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid14499155_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBöhmSchild2003" class="citation journal cs1">Böhm I, Schild H (2003). "Apoptosis: the complex scenario for a silent cell death". <i>Mol Imaging Biol</i>. <b>5</b> (1): 2–14. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS1536-1632%2803%2900024-6">10.1016/S1536-1632(03)00024-6</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14499155">14499155</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Mol+Imaging+Biol&rft.atitle=Apoptosis%3A+the+complex+scenario+for+a+silent+cell+death&rft.volume=5&rft.issue=1&rft.pages=2-14&rft.date=2003&rft_id=info%3Adoi%2F10.1016%2FS1536-1632%2803%2900024-6&rft_id=info%3Apmid%2F14499155&rft.aulast=B%C3%B6hm&rft.aufirst=I&rft.au=Schild%2C+H&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-FOOTNOTEAlberts2-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEAlberts2_3-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFAlberts">Alberts</a>, p. 2.</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKaram2009" class="citation book cs1">Karam JA (2009). <i>Apoptosis in Carcinogenesis and Chemotherapy</i>. Netherlands: Springer. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4020-9597-9" title="Special:BookSources/978-1-4020-9597-9"><bdi>978-1-4020-9597-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Apoptosis+in+Carcinogenesis+and+Chemotherapy&rft.place=Netherlands&rft.pub=Springer&rft.date=2009&rft.isbn=978-1-4020-9597-9&rft.aulast=Karam&rft.aufirst=JA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlbertsJohnsonLewisRaff2008" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2008). "Chapter 18 Apoptosis: Programmed Cell Death Eliminates Unwanted Cells". <a href="/wiki/Molecular_Biology_of_the_Cell_(textbook)" class="mw-redirect" title="Molecular Biology of the Cell (textbook)"><i>Molecular Biology of the Cell (textbook)</i></a> (5th ed.). <a href="/wiki/Garland_Science" title="Garland Science">Garland Science</a>. p. 1115. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-4105-5" title="Special:BookSources/978-0-8153-4105-5"><bdi>978-0-8153-4105-5</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+18+Apoptosis%3A+Programmed+Cell+Death+Eliminates+Unwanted+Cells&rft.btitle=Molecular+Biology+of+the+Cell+%28textbook%29&rft.pages=1115&rft.edition=5th&rft.pub=Garland+Science&rft.date=2008&rft.isbn=978-0-8153-4105-5&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%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRaychaudhuri2010" class="citation journal cs1">Raychaudhuri S (August 2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2920308">"A minimal model of signaling network elucidates cell-to-cell stochastic variability in apoptosis"</a>. <i>PLOS ONE</i>. <b>5</b> (8): e11930. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1009.2294">1009.2294</a></span>. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2010PLoSO...511930R">2010PLoSO...511930R</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0011930">10.1371/journal.pone.0011930</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/PMC2920308">2920308</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/20711445">20711445</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=A+minimal+model+of+signaling+network+elucidates+cell-to-cell+stochastic+variability+in+apoptosis&rft.volume=5&rft.issue=8&rft.pages=e11930&rft.date=2010-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2920308%23id-name%3DPMC&rft_id=info%3Abibcode%2F2010PLoSO...511930R&rft_id=info%3Aarxiv%2F1009.2294&rft_id=info%3Apmid%2F20711445&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0011930&rft.aulast=Raychaudhuri&rft.aufirst=S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2920308&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFElmore2007" class="citation journal cs1">Elmore S (June 2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2117903">"Apoptosis: A Review of Programmed Cell Death"</a>. <i>Toxicologic Pathology</i>. <b>35</b> (4): 495–516. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F01926230701320337">10.1080/01926230701320337</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/PMC2117903">2117903</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/17562483">17562483</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Toxicologic+Pathology&rft.atitle=Apoptosis%3A+A+Review+of+Programmed+Cell+Death&rft.volume=35&rft.issue=4&rft.pages=495-516&rft.date=2007-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2117903%23id-name%3DPMC&rft_id=info%3Apmid%2F17562483&rft_id=info%3Adoi%2F10.1080%2F01926230701320337&rft.aulast=Elmore&rft.aufirst=S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2117903&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-9">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKerr1965" class="citation journal cs1">Kerr JF (October 1965). "A histochemical study of hypertrophy and ischaemic injury of rat liver with special reference to changes in lysosomes". <i>The Journal of Pathology and Bacteriology</i>. <b>90</b> (2): 419–435. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fpath.1700900210">10.1002/path.1700900210</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/5849603">5849603</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Pathology+and+Bacteriology&rft.atitle=A+histochemical+study+of+hypertrophy+and+ischaemic+injury+of+rat+liver+with+special+reference+to+changes+in+lysosomes&rft.volume=90&rft.issue=2&rft.pages=419-435&rft.date=1965-10&rft_id=info%3Adoi%2F10.1002%2Fpath.1700900210&rft_id=info%3Apmid%2F5849603&rft.aulast=Kerr&rft.aufirst=JF&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-AHW_1972-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-AHW_1972_10-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAgency_for_Science,_Technology_and_Research" class="citation web cs1">Agency for Science, Technology and Research. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20071113101931/http://www.a-star.edu.sg/astar/biomed/action/biomed_dvp_abstract.do?id=2901ddeb02dH">"Prof Andrew H. Wyllie – Lecture Abstract"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.a-star.edu.sg/astar/biomed/action/biomed_dvp_abstract.do?id=2901ddeb02dH">the original</a> on 2007-11-13<span class="reference-accessdate">. Retrieved <span class="nowrap">2007-03-30</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Prof+Andrew+H.+Wyllie+%E2%80%93+Lecture+Abstract&rft.au=Agency+for+Science%2C+Technology+and+Research&rft_id=http%3A%2F%2Fwww.a-star.edu.sg%2Fastar%2Fbiomed%2Faction%2Fbiomed_dvp_abstract.do%3Fid%3D2901ddeb02dH&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-Kerr1972-11"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kerr1972_11-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kerr1972_11-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKerrWyllieCurrie1972" class="citation journal cs1">Kerr JF, Wyllie AH, Currie AR (August 1972). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2008650">"Apoptosis: a basic biological phenomenon with wide-ranging implications in tissue kinetics"</a>. <i>British Journal of Cancer</i>. <b>26</b> (4): 239–257. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fbjc.1972.33">10.1038/bjc.1972.33</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/PMC2008650">2008650</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/4561027">4561027</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=British+Journal+of+Cancer&rft.atitle=Apoptosis%3A+a+basic+biological+phenomenon+with+wide-ranging+implications+in+tissue+kinetics&rft.volume=26&rft.issue=4&rft.pages=239-257&rft.date=1972-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2008650%23id-name%3DPMC&rft_id=info%3Apmid%2F4561027&rft_id=info%3Adoi%2F10.1038%2Fbjc.1972.33&rft.aulast=Kerr&rft.aufirst=JF&rft.au=Wyllie%2C+AH&rft.au=Currie%2C+AR&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2008650&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFO'RourkeEllem2000" class="citation journal cs1">O'Rourke MG, Ellem KA (2000). 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"Bcl-2 gene promotes haemopoietic cell survival and cooperates with c-myc to immortalize pre-B cells". <i>Nature</i>. <b>335</b> (6189): 440–2. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1988Natur.335..440V">1988Natur.335..440V</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F335440a0">10.1038/335440a0</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/3262202">3262202</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:23593952">23593952</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Bcl-2+gene+promotes+haemopoietic+cell+survival+and+cooperates+with+c-myc+to+immortalize+pre-B+cells&rft.volume=335&rft.issue=6189&rft.pages=440-2&rft.date=1988-09&rft_id=info%3Adoi%2F10.1038%2F335440a0&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A23593952%23id-name%3DS2CID&rft_id=info%3Apmid%2F3262202&rft_id=info%3Abibcode%2F1988Natur.335..440V&rft.aulast=Vaux&rft.aufirst=DL&rft.au=Cory%2C+S&rft.au=Adams%2C+JM&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> <li id="cite_note-FOOTNOTEAlberts1021-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEAlberts1021_14-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFAlberts">Alberts</a>, p. 1021.</span> </li> <li id="cite_note-ahdictionary-15"><span class="mw-cite-backlink">^ <a href="#cite_ref-ahdictionary_15-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-ahdictionary_15-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://ahdictionary.com/word/search.html?q=apoptosis">"The American Heritage Dictionary entry: apoptosis"</a>. <i>ahdictionary.com</i>. Houghton Mifflin Harcourt Publishing Company. 2020. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210726211047/https://ahdictionary.com/word/search.html?q=apoptosis">Archived</a> from the original on 26 July 2021<span class="reference-accessdate">. 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"Atromentin-induced apoptosis in human leukemia U937 cells". <i>Journal of Microbiology and Biotechnology</i>. <b>19</b> (9): 946–950. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.4014%2Fjmb.0811.617">10.4014/jmb.0811.617</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19809251">19809251</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:11552839">11552839</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Microbiology+and+Biotechnology&rft.atitle=Atromentin-induced+apoptosis+in+human+leukemia+U937+cells&rft.volume=19&rft.issue=9&rft.pages=946-950&rft.date=2009-09&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A11552839%23id-name%3DS2CID&rft_id=info%3Apmid%2F19809251&rft_id=info%3Adoi%2F10.4014%2Fjmb.0811.617&rft.aulast=Kim&rft.aufirst=JH&rft.au=Lee%2C+CH&rfr_id=info%3Asid%2Fen.wikipedia.org%3AApoptosis" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="General_bibliography">General bibliography</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Apoptosis&action=edit&section=32" title="Edit section: General bibliography"><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" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlberts" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2015). <i>Molecular Biology of the Cell</i> (6th ed.). Garland Science. p. 2. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0815344322" title="Special:BookSources/978-0815344322"><bdi>978-0815344322</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+Biology+of+the+Cell&rft.pages=2&rft.edition=6th&rft.pub=Garland+Science&rft.date=2015&rft.isbn=978-0815344322&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%3AApoptosis" 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=Apoptosis&action=edit&section=33" 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><div class="side-box metadata side-box-right"><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"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Scholia_logo.svg/40px-Scholia_logo.svg.png" decoding="async" width="40" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Scholia_logo.svg/60px-Scholia_logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/Scholia_logo.svg/80px-Scholia_logo.svg.png 2x" data-file-width="107" data-file-height="104" /></span></span></div> <div class="side-box-text plainlist"><a href="https://www.wikidata.org/wiki/Wikidata:Scholia" class="extiw" title="d:Wikidata:Scholia">Scholia</a> has a profile for <a href="https://iw.toolforge.org/scholia/Q29892216" class="extiw" title="toolforge:scholia/Q29892216"><b>Apoptosis <small>(Q29892216)</small></b></a>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=l4D0YxGi5Ec">Apoptosis & Caspase 3</a>, <a href="/wiki/The_Proteolysis_Map" title="The Proteolysis Map">The Proteolysis Map</a> – animation</li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=29AMumxsEo0">Apoptosis & Caspase 8</a>, <a href="/wiki/The_Proteolysis_Map" title="The Proteolysis Map">The Proteolysis Map</a> – animation</li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=4YYboqiol_w">Apoptosis & Caspase 7</a>, <a href="/wiki/The_Proteolysis_Map" title="The Proteolysis Map">The Proteolysis Map</a> – animation</li> <li><a rel="nofollow" class="external text" href="http://www.copewithcytokines.de/cope.cgi?key=Apoptosis%20MiniCOPE%20Dictionary">Apoptosis MiniCOPE Dictionary – list of apoptosis terms and acronyms</a></li> <li><a rel="nofollow" class="external text" href="http://www.biochemweb.net/apoptosis.shtml">Apoptosis (Programmed Cell Death) – The Virtual Library of Biochemistry, Molecular Biology and Cell Biology</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210425180849/http://www.biochemweb.net/apoptosis.shtml">Archived</a> 2021-04-25 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20140825065034/http://www.caspases.org/">Apoptosis Research Portal</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070111132934/http://apoptosisinfo.com/">Apoptosis Info</a> Apoptosis protocols, articles, news, and recent publications.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070111190837/http://www.apoptosis-db.org/welcome.html">Database of proteins involved in apoptosis</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070630103651/http://stke.sciencemag.org/content/vol2007/issue380/images/data/tr1/DC1/Apoptosis_WEHI.mov">Apoptosis Video</a></li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=DR80Huxp4y8">Apoptosis Video (WEHI on YouTube )</a></li> <li><a rel="nofollow" class="external text" href="http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/A/Apoptosis.html#The_Mechanisms_of_Apoptosis">The Mechanisms of Apoptosis</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180309073759/http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/A/Apoptosis.html#The_Mechanisms_of_Apoptosis">Archived</a> 2018-03-09 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> Kimball's Biology Pages. Simple explanation of the mechanisms of apoptosis triggered by internal signals (bcl-2), along the caspase-9, caspase-3 and caspase-7 pathway; and by external signals (FAS and TNF), along the caspase 8 pathway. Accessed 25 March 2007.</li> <li>WikiPathways – <a rel="nofollow" class="external text" href="http://www.wikipathways.org/index.php/Pathway:Homo_sapiens:Apoptosis">Apoptosis pathway</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080916135457/http://www.wikipathways.org/index.php/Pathway:Homo_sapiens:Apoptosis">Archived</a> 2008-09-16 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20141227072014/http://www.crmagazine.org/archive/Spring2007/Pages/FindingCancersSelf-DestructButton.aspx">"Finding Cancer's Self-Destruct Button"</a>. <i>CR</i> magazine (Spring 2007). Article on apoptosis and cancer.</li> <li><a rel="nofollow" class="external text" href="http://www.ibiology.org/ibioseminars/cell-biology/xiaodong-wang-part-1.html">Xiaodong Wang's lecture: Introduction to Apoptosis</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20131029222808/http://www.ibiology.org/ibioseminars/cell-biology/xiaodong-wang-part-1.html">Archived</a> 2013-10-29 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://www.ibiology.org/cell-biology/programmed-cell-death/">Robert Horvitz's Short Clip: Discovering Programmed Cell Death</a></li> <li><a rel="nofollow" class="external text" href="http://bcl2db.ibcp.fr/BCL2DB/">The Bcl-2 Database</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20131023061339/http://bcl2db.ibcp.fr/BCL2DB/">Archived</a> 2013-10-23 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="http://arquivo.pt/wayback/20160515224643/http://www.deathbase.org/">DeathBase: a database of proteins involved in cell death, curated by experts</a></li> <li><a rel="nofollow" class="external text" href="http://www.ecdo.eu">European Cell Death Organization</a></li> <li><a rel="nofollow" class="external text" href="https://www.cusabio.com/Apoptosis/">Apoptosis signaling pathway</a> created by Cusabio</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl 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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_proteins" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_cycle" title="Cell cycle">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 href="/wiki/Cell_cycle#Inhibitors" title="Cell cycle">cip/kip</a> (<a href="/wiki/P21" title="P21">p21</a>, <a href="/wiki/CDKN1B" title="CDKN1B">p27</a>, <a href="/wiki/Cyclin-dependent_kinase_inhibitor_1C" title="Cyclin-dependent kinase inhibitor 1C">p57</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/P53_p63_p73_family" title="P53 p63 p73 family">P53 p63 p73 family</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/P53" title="P53">p53</a></li> <li><a href="/wiki/TP63" title="TP63">p63</a></li> <li><a href="/wiki/P73" title="P73">p73</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cdk1" class="mw-redirect" title="Cdk1">Cdc2</a></li> <li><a href="/wiki/Cdc25" title="Cdc25">Cdc25</a></li> <li><a href="/wiki/CDC42" title="CDC42">Cdc42</a></li> <li><a href="/wiki/Cellular_apoptosis_susceptibility_protein" title="Cellular apoptosis susceptibility protein">Cellular apoptosis susceptibility protein</a></li> <li><a href="/wiki/E2F" title="E2F">E2F</a></li> <li><a href="/wiki/Maturation_promoting_factor" title="Maturation promoting factor">Maturation promoting factor</a></li> <li><a href="/wiki/Wee1" title="Wee1">Wee</a></li> <li><a href="/wiki/Cullin" title="Cullin">Cullin</a> (<a href="/wiki/CUL7" title="CUL7">CUL7</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Phases and<br />checkpoints</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Interphase" title="Interphase">Interphase</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/G1_phase" title="G1 phase">G<sub>1</sub> phase</a></li> <li><a href="/wiki/S_phase" title="S phase">S phase</a></li> <li><a href="/wiki/G2_phase" title="G2 phase">G<sub>2</sub> phase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cell_division" title="Cell division">M phase</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitosis" title="Mitosis">Mitosis</a> (<a href="/wiki/Preprophase" title="Preprophase">Preprophase</a></li> <li><a href="/wiki/Prophase" title="Prophase">Prophase</a></li> <li><a href="/wiki/Prometaphase" title="Prometaphase">Prometaphase</a></li> <li><a href="/wiki/Metaphase" title="Metaphase">Metaphase</a></li> <li><a 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 class="mw-selflink selflink">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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Cell_signaling_/_Signal_transduction" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Signal_transduction" title="Template:Signal transduction"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Signal_transduction" title="Template talk:Signal transduction"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Signal_transduction" title="Special:EditPage/Template:Signal transduction"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_signaling_/_Signal_transduction" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_signaling" title="Cell signaling">Cell signaling</a> / <a href="/wiki/Signal_transduction" title="Signal transduction">Signal transduction</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Signaling pathways</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/G_protein-coupled_receptor" title="G protein-coupled receptor">GPCR</a></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt</a></li> <li><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">RTK</a> <ul><li><a href="/wiki/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta</a></li> <li><a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK</a></li></ul></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch</a></li> <li><a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT</a></li> <li><a href="/wiki/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB</a></li> <li><a class="mw-selflink selflink">Fas apoptosis</a></li> <li><a href="/wiki/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo</a></li> <li><a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a></li> <li><a href="/wiki/Integrin#Signal_transduction" title="Integrin">Integrin receptors</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Agents</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/Ligand_(biochemistry)" title="Ligand (biochemistry)">Receptor ligands</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/Hormone" title="Hormone">Hormones</a></li> <li><a href="/wiki/Neurotransmitter" title="Neurotransmitter">Neurotransmitters</a>/<a href="/wiki/Neuropeptide" title="Neuropeptide">Neuropeptides</a>/<a href="/wiki/Neurohormone" title="Neurohormone">Neurohormones</a></li> <li><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a></li> <li><a href="/wiki/Growth_factor" title="Growth factor">Growth factors</a></li> <li><a href="/wiki/Cell_signaling" title="Cell signaling">Signaling molecules</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">Cell surface</a></li> <li><a href="/wiki/Intracellular_receptor" title="Intracellular receptor">Intracellular</a></li> <li><a href="/wiki/Co-receptor" title="Co-receptor">Co-receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Second_messenger_system" title="Second messenger system">Second messenger</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP-dependent pathway</a></li> <li><a href="/wiki/Calcium_signaling" title="Calcium signaling">Ca<sup>2+</sup> signaling</a></li> <li><a href="/wiki/Lipid_signaling" title="Lipid signaling">Lipid signaling</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Assistants:</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/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">Signal transducing adaptor protein</a></li> <li><a href="/wiki/Scaffold_protein" title="Scaffold protein">Scaffold protein</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factors</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/General_transcription_factor" title="General transcription factor">General</a></li> <li><a href="/wiki/Transcription_preinitiation_complex" title="Transcription preinitiation complex">Transcription preinitiation complex</a></li> <li><a href="/wiki/Transcription_factor_II_D" title="Transcription factor II D">TFIID</a></li> <li><a href="/wiki/Transcription_factor_II_H" title="Transcription factor II H">TFIIH</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By distance</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/Juxtacrine_signalling" title="Juxtacrine signalling">Juxtacrine</a></li> <li><a href="/wiki/Autocrine_signalling" class="mw-redirect" title="Autocrine signalling">Autocrine</a> / <a href="/wiki/Paracrine_signalling" class="mw-redirect" title="Paracrine signalling">Paracrine</a></li> <li><a href="/wiki/Endocrine_system" title="Endocrine system">Endocrine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other concepts</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/Intracrine" title="Intracrine">Intracrine action</a></li> <li>Neurocrine signaling <ul><li><a href="/wiki/Synaptic_transmission" class="mw-redirect" title="Synaptic transmission">Synaptic transmission</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a></li></ul></li> <li><a href="/wiki/Neuroendocrine_cell" title="Neuroendocrine cell">Neuroendocrine signaling</a></li> <li><a href="/wiki/Exocrine_gland" title="Exocrine gland">Exocrine signalling</a> <ul><li><a href="/wiki/Pheromone" title="Pheromone">Pheromones</a></li></ul></li> <li><a href="/wiki/Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a></li> <li><a href="/wiki/Visual_phototransduction" title="Visual phototransduction">Phototransduction</a></li> <li><a href="/wiki/Ion_channel" title="Ion channel">Ion channel gating</a></li> <li><a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Apoptosis_signaling_pathway" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:#e7dcc3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Apoptosis_signaling_pathway" title="Template:Apoptosis signaling pathway"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Apoptosis_signaling_pathway" title="Template talk:Apoptosis signaling pathway"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Apoptosis_signaling_pathway" title="Special:EditPage/Template:Apoptosis signaling pathway"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Apoptosis_signaling_pathway" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Apoptosis</a> <a href="/wiki/Signal_transduction" title="Signal transduction">signaling pathway</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink-fragment" href="#Fas_path">Fas path</a></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%;background-color: AntiqueWhite"><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</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/Fas_ligand" title="Fas ligand">Fas ligand</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptor</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/Fas_receptor" title="Fas receptor">Fas receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Death-inducing_signaling_complex" title="Death-inducing signaling complex">Death-inducing signaling complex</a></li></ul> <ul><li><a href="/wiki/DAXX" class="mw-redirect" title="DAXX">DAXX</a></li> <li><a href="/wiki/ASK1" title="ASK1">ASK1</a></li></ul> <ul><li><a href="/wiki/FADD" title="FADD">FADD</a></li> <li><a href="/wiki/Caspase_8" title="Caspase 8">Caspase 8</a></li> <li><a href="/wiki/BH3_interacting_domain_death_agonist" class="mw-redirect" title="BH3 interacting domain death agonist">BID</a></li></ul> <ul><li><a href="/wiki/Cytochrome_c" title="Cytochrome c">Cytochrome c</a></li> <li><a href="/wiki/Caspase-9" title="Caspase-9">Caspase 9</a></li> <li><a href="/wiki/Caspase_3" title="Caspase 3">Caspase 3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Bcl-2_family" title="Bcl-2 family">Bcl-2 family</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><i>Pro-apoptotic:</i></dt> <dd><a href="/wiki/Bcl-2-associated_X_protein" class="mw-redirect" title="Bcl-2-associated X protein">BAX</a></dd> <dd>BAK1/<a href="/wiki/Bcl-2_homologous_antagonist_killer" title="Bcl-2 homologous antagonist killer">Bcl-2 homologous antagonist killer</a></dd> <dd><a href="/wiki/Bcl-2-associated_death_promoter" title="Bcl-2-associated death promoter">Bcl-2-associated death promoter</a></dd></dl> <dl><dt><i>Anti-apoptotic:</i></dt> <dd><a href="/wiki/Bcl-2" title="Bcl-2">Bcl-2</a></dd> <dd><a href="/wiki/Bcl-xL" title="Bcl-xL">Bcl-xL</a></dd></dl> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink-fragment" href="#TNF_path">TNF path</a></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%;background-color: AntiqueWhite"><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</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/Tumor_necrosis_factor_alpha" class="mw-redirect" title="Tumor necrosis factor alpha">Tumor necrosis factor alpha</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptor</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/Tumor_necrosis_factor_receptor_1" title="Tumor necrosis factor receptor 1">Tumor necrosis factor receptor 1</a></li> <li><a href="/wiki/Tumor_necrosis_factor_receptor_2" title="Tumor necrosis factor receptor 2">Tumor necrosis factor receptor 2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular</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/TRADD" title="TRADD">TRADD</a></li></ul> <ul><li><a href="/wiki/FADD" title="FADD">FADD</a></li> <li><a href="/wiki/Caspase_8" title="Caspase 8">Caspase 8</a></li> <li><a href="/wiki/Caspase_3" title="Caspase 3">Caspase 3</a></li> <li><a href="/wiki/BH3_interacting_domain_death_agonist" class="mw-redirect" title="BH3 interacting domain death agonist">BID</a></li></ul> <ul><li><a href="/wiki/TRAF2" title="TRAF2">TRAF2</a></li> <li><a href="/wiki/ASK1" title="ASK1">ASK-1</a></li> <li><a href="/wiki/MAP3K1" title="MAP3K1">MEKK1</a></li> <li><a href="/wiki/I%CE%BAB_kinase" title="IκB kinase">IKK</a></li> <li><a href="/wiki/I%CE%BAB%CE%B1" title="IκBα">IκBα</a></li> <li><a href="/wiki/MAP2K7" title="MAP2K7">MKK7</a></li> <li><a href="/wiki/C-Jun_N-terminal_kinases" title="C-Jun N-terminal kinases">JNK</a></li> <li><a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-κB</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other</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 id="Intracellular" scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite; text-align:center; padding-left:3.6em;"><a href="/wiki/Intracellular_signaling_peptides_and_proteins" class="mw-redirect" title="Intracellular signaling peptides and proteins">Intracellular</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">IAPs</a></dt> <dd><a href="/wiki/XIAP" title="XIAP">XIAP</a></dd> <dd><a href="/wiki/NAIP_(gene)" title="NAIP (gene)">NAIP</a></dd> <dd><a href="/wiki/Survivin" title="Survivin">Survivin</a></dd> <dd><a href="/wiki/BIRC2" class="mw-redirect" title="BIRC2">c-IAP-1</a></dd> <dd><a href="/wiki/BIRC3" class="mw-redirect" title="BIRC3">c-IAP-2</a></dd></dl> <ul><li><a href="/wiki/Apoptosis-inducing_factor" title="Apoptosis-inducing factor">Apoptosis-inducing factor</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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q14599311#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q14599311#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q14599311#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Apoptosis"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh91000743">United States</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Apoptose"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb12463578d">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Apoptose"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb12463578d">BnF data</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/01046102">Japan</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="apoptóza"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph182376&CON_LNG=ENG">Czech Republic</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007548987205171">Israel</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="http://esu.com.ua/search_articles.php?id=43113">Encyclopedia of Modern Ukraine</a></span></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&useformat=desktop" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" 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12 Template:Cn"," 6.63% 127.884 6 Template:Sfn"," 6.19% 119.411 1 Template:Langx"," 5.83% 112.499 10 Template:Navbox"]},"scribunto":{"limitreport-timeusage":{"value":"1.263","limit":"10.000"},"limitreport-memusage":{"value":20793836,"limit":52428800},"limitreport-logs":"anchor_id_list = table#1 {\n [\"CITEREFAcraniGomesProença-Módenada_Silva2010\"] = 1,\n [\"CITEREFAgency_for_Science,_Technology_and_Research\"] = 1,\n [\"CITEREFAlberts\"] = 1,\n [\"CITEREFAlbertsJohnsonLewisRaff2008\"] = 1,\n [\"CITEREFAlimontiBallFowke2003\"] = 1,\n [\"CITEREFApoptosis_Interest_Group1999\"] = 1,\n [\"CITEREFAtkin-SmithTixeiraPaoneMathivanan2015\"] = 1,\n [\"CITEREFAzevedoNunesChiangBensabath2007\"] = 1,\n [\"CITEREFBejaranoEspinoGonzález-FloresCasado2009\"] = 1,\n [\"CITEREFBernsteinBernsteinPayneGarewal2002\"] = 1,\n [\"CITEREFBoehm2006\"] = 1,\n [\"CITEREFBrattonSalvesen2010\"] = 1,\n [\"CITEREFBrüne2003\"] = 1,\n [\"CITEREFBrünevon_KnethenSandau1999\"] = 1,\n [\"CITEREFBöhm2003\"] = 1,\n [\"CITEREFBöhmSchild2003\"] = 1,\n [\"CITEREFChenGoeddel2002\"] = 1,\n [\"CITEREFChenLiChenHan2005\"] = 1,\n [\"CITEREFChiarugiMoskowitz2002\"] = 1,\n [\"CITEREFColemanSahaiYeoBosch2001\"] = 1,\n [\"CITEREFCollazoChacónBorrás2006\"] = 1,\n [\"CITEREFCotranKumar1998\"] = 1,\n [\"CITEREFDarzynkiewiczJuanLiGorczyca1997\"] = 1,\n [\"CITEREFDel_PuertoMartinsMilstedSouza-Fagundes2011\"] = 1,\n [\"CITEREFDemetriusMagistrettiPellerin2014\"] = 1,\n [\"CITEREFDickmanWilliamsLide_Figueiredo2017\"] = 1,\n [\"CITEREFElmore2007\"] = 1,\n [\"CITEREFEverettMcFadden1999\"] = 1,\n [\"CITEREFFarhanaDawsonFontana2005\"] = 1,\n [\"CITEREFFesikShi2001\"] = 1,\n [\"CITEREFGerlVaux2005\"] = 1,\n [\"CITEREFGoeddelDV2007\"] = 1,\n [\"CITEREFGoldsteinWaterhouseJuinEvan2000\"] = 1,\n [\"CITEREFGongTraganosDarzynkiewicz1994\"] = 1,\n [\"CITEREFGonzalezBejaranoBarrigaRodriguez2010\"] = 1,\n [\"CITEREFGreen2011\"] = 1,\n [\"CITEREFHalickaBednerDarzynkiewicz2000\"] = 1,\n [\"CITEREFHardyEl-AssaadPrzybytkowskiJoly2003\"] = 1,\n [\"CITEREFHayKannourakis2002\"] = 1,\n [\"CITEREFIndiana_University_Health\"] = 1,\n [\"CITEREFIndranTufoPervaizBrenner2011\"] = 1,\n [\"CITEREFIwataMyersonTorok-StorbZager1994\"] = 1,\n [\"CITEREFJanChaudhry2019\"] = 1,\n [\"CITEREFJewhurstLevinMcLaughlin2014\"] = 1,\n [\"CITEREFKaczanowski2016\"] = 1,\n [\"CITEREFKaleOsterlundAndrews2018\"] = 1,\n [\"CITEREFKaram2009\"] = 1,\n [\"CITEREFKerr1965\"] = 1,\n [\"CITEREFKerrWyllieCurrie1972\"] = 1,\n [\"CITEREFKihlmarkImrehHallberg2001\"] = 1,\n [\"CITEREFKimLee2009\"] = 1,\n [\"CITEREFKryskoVanden_BergheD\u0026#039;HerdeVandenabeele2008\"] = 1,\n [\"CITEREFKryskoVanden_BergheParthoensD\u0026#039;Herde2008\"] = 1,\n [\"CITEREFKryskoVandenabeele2009\"] = 1,\n [\"CITEREFLaFerlaTinkleBieberichHaudenschild1995\"] = 1,\n [\"CITEREFLeeLuMadhukar2010\"] = 1,\n [\"CITEREFLiSarkisianMehalRakic2003\"] = 1,\n [\"CITEREFLiuChenVlantisTse2008\"] = 1,\n [\"CITEREFLiuMcKalipHerman2000\"] = 1,\n [\"CITEREFLozanoBejaranoEspinoGonzález2009\"] = 1,\n [\"CITEREFMailhosHowardLatchman1994\"] = 2,\n [\"CITEREFMarinoDunnGrailInglese1997\"] = 1,\n [\"CITEREFMasumYokoiHisamatsuNaito2018\"] = 1,\n [\"CITEREFMattsonChan2003\"] = 1,\n [\"CITEREFMochizukiGotoMoriMizuno1996\"] = 1,\n [\"CITEREFMossBetinMalesinskiLane2006\"] = 1,\n [\"CITEREFMurphyRanganathanFarnsworthKavallaris2000\"] = 1,\n [\"CITEREFMusiccoAdorniDi_SantoPrinelli2013\"] = 1,\n [\"CITEREFNagata2000\"] = 1,\n [\"CITEREFNirmalaLopus2020\"] = 1,\n [\"CITEREFNishiTsukiyama-KoharaTogashiKohriyama2004\"] = 1,\n [\"CITEREFNiuPengDuanWang2006\"] = 1,\n [\"CITEREFO\u0026#039;RourkeEllem2000\"] = 1,\n [\"CITEREFPolsterPevsnerHardwick2004\"] = 1,\n [\"CITEREFPoonChiuArmstrongKinchen2014\"] = 1,\n [\"CITEREFRaychaudhuri2010\"] = 1,\n [\"CITEREFRazaghiHeimannSchaefferGibson2018\"] = 1,\n [\"CITEREFSantosRodriguesSilvaMortara2008\"] = 1,\n [\"CITEREFSavillGregoryHaslett2003\"] = 1,\n [\"CITEREFSebbaghRenvoizéHamelinRiché2001\"] = 1,\n [\"CITEREFSmithParkesAtkin-SmithTixeira2017\"] = 1,\n [\"CITEREFSusinDaugasRavagnanSamejima2000\"] = 1,\n [\"CITEREFSusinLorenzoZamzamiMarzo1999\"] = 1,\n [\"CITEREFTakaokaHayakawaYanaiStoiber2003\"] = 1,\n [\"CITEREFTamuraTakayamaIshiiMawaribuchi2015\"] = 1,\n [\"CITEREFTateishiMondeAnrakuKoga2017\"] = 1,\n [\"CITEREFTeodoroBranton1997\"] = 1,\n [\"CITEREFThomasLiuWhangboMcCrossan2015\"] = 1,\n [\"CITEREFThompson1995\"] = 1,\n [\"CITEREFTixeiraCarusoPaoneBaxter2017\"] = 1,\n [\"CITEREFUrenIyerKluck2017\"] = 1,\n [\"CITEREFUğuzNaziroğluEspinoBejarano2009\"] = 1,\n [\"CITEREFVanden_BergheGrootjansGoossensDondelinger2013\"] = 1,\n [\"CITEREFVandivierHensonDouglas2006\"] = 1,\n [\"CITEREFVashisthaHusainKumarYadav2008\"] = 1,\n [\"CITEREFVauxCoryAdams1988\"] = 1,\n [\"CITEREFVenturi,_Sebastiano2014\"] = 1,\n [\"CITEREFVenturi2011\"] = 1,\n [\"CITEREFVlahopoulos2017\"] = 1,\n [\"CITEREFWajant2002\"] = 1,\n [\"CITEREFWajant2007\"] = 1,\n [\"CITEREFWangGibsonVermeulenYeh1995\"] = 1,\n [\"CITEREFWangWuFadokLee2003\"] = 1,\n [\"CITEREFWangYoule2009\"] = 1,\n [\"CITEREFWarburg1956\"] = 1,\n [\"CITEREFWestphalKluckDewson2014\"] = 1,\n [\"CITEREFWlodkowicTelfordSkommerDarzynkiewicz2011\"] = 1,\n [\"CITEREFXiangChaoKorsmeyer1996\"] = 1,\n [\"CITEREFYangMashimaSatoMochizuki2003\"] = 1,\n [\"CITEREFvan_DoornBeersDanglFranklin-Tong2011\"] = 1,\n}\ntemplate_list = table#1 {\n [\"Apoptosis signaling pathway\"] = 1,\n [\"Authority control\"] = 1,\n [\"Better source\"] = 1,\n [\"Cell cycle proteins\"] = 1,\n [\"Citation needed\"] = 6,\n [\"Cite book\"] = 8,\n [\"Cite journal\"] = 100,\n [\"Cite web\"] = 5,\n [\"Cn\"] = 12,\n [\"Div col\"] = 1,\n [\"Div col end\"] = 1,\n [\"Efn\"] = 2,\n [\"Further\"] = 1,\n [\"IPAc-en\"] = 2,\n [\"Infobox anatomy\"] = 1,\n [\"Lang\"] = 1,\n [\"Langx\"] = 1,\n [\"Main\"] = 2,\n [\"Multiple image\"] = 1,\n [\"Notelist\"] = 1,\n [\"Portal\"] = 1,\n [\"Refbegin\"] = 1,\n [\"Refend\"] = 1,\n [\"Reflist\"] = 1,\n [\"Respell\"] = 1,\n [\"Retracted\"] = 1,\n [\"Scholia\"] = 1,\n [\"Sfn\"] = 6,\n [\"SfnRef\"] = 1,\n [\"Short description\"] = 1,\n [\"Signal transduction\"] = 1,\n [\"Val\"] = 2,\n [\"Webarchive\"] = 5,\n}\narticle_whitelist = table#1 {\n}\n","limitreport-profile":[["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::callParserFunction","300","23.1"],["?","180","13.8"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::match","80","6.2"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::getAllExpandedArguments","80","6.2"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::gsub","80","6.2"],["dataWrapper \u003Cmw.lua:672\u003E","60","4.6"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::sub","60","4.6"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::getEntity","40","3.1"],["newFrame \u003Cmw.lua:153\u003E","40","3.1"],["validateData \u003Cmw.lua:728\u003E","40","3.1"],["[others]","340","26.2"]]},"cachereport":{"origin":"mw-web.codfw.main-5c59558b9d-l5884","timestamp":"20241130110549","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Apoptosis","url":"https:\/\/en.wikipedia.org\/wiki\/Apoptosis","sameAs":"http:\/\/www.wikidata.org\/entity\/Q14599311","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q14599311","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2001-11-05T16:59:01Z","dateModified":"2024-11-29T14:25:32Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/2\/23\/Apoptosis_DU145_cells_mosaic.jpg","headline":"programmed cell death in multicellular organisms"}</script> </body> 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