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</li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> 2 Function </div> </a> <button aria-controls="toc-Function-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Function subsection </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Energy_conversion" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Energy_conversion"> <div class="vector-toc-text"> 2.1 Energy conversion </div> </a> <ul id="toc-Energy_conversion-sublist" class="vector-toc-list"> <li id="toc-Pyruvate_and_the_citric_acid_cycle" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Pyruvate_and_the_citric_acid_cycle"> <div class="vector-toc-text"> 2.1.1 Pyruvate and the citric acid cycle </div> </a> <ul id="toc-Pyruvate_and_the_citric_acid_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-O2_and_NADH:_energy-releasing_reactions" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#O2_and_NADH:_energy-releasing_reactions"> <div class="vector-toc-text"> 2.1.2 O2 and NADH: energy-releasing reactions </div> </a> <ul id="toc-O2_and_NADH:_energy-releasing_reactions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Heat_production" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Heat_production"> <div class="vector-toc-text"> 2.1.3 Heat production </div> </a> <ul id="toc-Heat_production-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mitochondrial_fatty_acid_synthesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitochondrial_fatty_acid_synthesis"> <div class="vector-toc-text"> 2.2 Mitochondrial fatty acid synthesis </div> </a> <ul id="toc-Mitochondrial_fatty_acid_synthesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Uptake,_storage_and_release_of_calcium_ions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Uptake,_storage_and_release_of_calcium_ions"> <div class="vector-toc-text"> 2.3 Uptake, storage and release of calcium ions </div> </a> <ul id="toc-Uptake,_storage_and_release_of_calcium_ions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cellular_proliferation_regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cellular_proliferation_regulation"> <div class="vector-toc-text"> 2.4 Cellular proliferation regulation </div> </a> <ul id="toc-Cellular_proliferation_regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Programmed_cell_death_and_innate_immunity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Programmed_cell_death_and_innate_immunity"> <div class="vector-toc-text"> 2.5 Programmed cell death and innate immunity </div> </a> <ul id="toc-Programmed_cell_death_and_innate_immunity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Additional_functions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Additional_functions"> <div class="vector-toc-text"> 2.6 Additional functions </div> </a> <ul id="toc-Additional_functions-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Organization_and_distribution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Organization_and_distribution"> <div class="vector-toc-text"> 3 Organization and distribution </div> </a> <button aria-controls="toc-Organization_and_distribution-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Organization and distribution subsection </button> <ul id="toc-Organization_and_distribution-sublist" class="vector-toc-list"> <li id="toc-Mitochondria-associated_ER_membrane_(MAM)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitochondria-associated_ER_membrane_(MAM)"> <div class="vector-toc-text"> 3.1 Mitochondria-associated ER membrane (MAM) </div> </a> <ul id="toc-Mitochondria-associated_ER_membrane_(MAM)-sublist" class="vector-toc-list"> <li id="toc-Phospholipid_transfer" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Phospholipid_transfer"> <div class="vector-toc-text"> 3.1.1 Phospholipid transfer </div> </a> <ul id="toc-Phospholipid_transfer-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Calcium_signaling" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Calcium_signaling"> <div class="vector-toc-text"> 3.1.2 Calcium signaling </div> </a> <ul id="toc-Calcium_signaling-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Molecular_basis_for_tethering" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Molecular_basis_for_tethering"> <div class="vector-toc-text"> 3.1.3 Molecular basis for tethering </div> </a> <ul id="toc-Molecular_basis_for_tethering-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Perspective" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Perspective"> <div class="vector-toc-text"> 3.1.4 Perspective </div> </a> <ul id="toc-Perspective-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Origin_and_evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Origin_and_evolution"> <div class="vector-toc-text"> 4 Origin and evolution </div> </a> <ul id="toc-Origin_and_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mitochondrial_genetics" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mitochondrial_genetics"> <div class="vector-toc-text"> 5 Mitochondrial genetics </div> </a> <button aria-controls="toc-Mitochondrial_genetics-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Mitochondrial genetics subsection </button> <ul id="toc-Mitochondrial_genetics-sublist" class="vector-toc-list"> <li id="toc-Human_population_genetic_studies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Human_population_genetic_studies"> <div class="vector-toc-text"> 5.1 Human population genetic studies </div> </a> <ul id="toc-Human_population_genetic_studies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_genetic_code" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Alternative_genetic_code"> <div class="vector-toc-text"> 5.2 Alternative genetic code </div> </a> <ul id="toc-Alternative_genetic_code-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Replication_and_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Replication_and_inheritance"> <div class="vector-toc-text"> 5.3 Replication and inheritance </div> </a> <ul id="toc-Replication_and_inheritance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_repair" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_repair"> <div class="vector-toc-text"> 5.4 DNA repair </div> </a> <ul id="toc-DNA_repair-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Lack_of_mitochondrial_DNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lack_of_mitochondrial_DNA"> <div class="vector-toc-text"> 5.5 Lack of mitochondrial DNA </div> </a> <ul id="toc-Lack_of_mitochondrial_DNA-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Dysfunction_and_disease" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Dysfunction_and_disease"> <div class="vector-toc-text"> 6 Dysfunction and disease </div> </a> <button aria-controls="toc-Dysfunction_and_disease-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Dysfunction and disease subsection </button> <ul id="toc-Dysfunction_and_disease-sublist" class="vector-toc-list"> <li id="toc-Mitochondrial_diseases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitochondrial_diseases"> <div class="vector-toc-text"> 6.1 Mitochondrial diseases </div> </a> <ul id="toc-Mitochondrial_diseases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Relationships_to_aging" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Relationships_to_aging"> <div class="vector-toc-text"> 6.2 Relationships to aging </div> </a> <ul id="toc-Relationships_to_aging-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> 7 History </div> </a> <ul id="toc-History-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"> 8 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 9 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 10 External links </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" > Toggle the table of contents </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">Mitochondrion</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 102 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-102" aria-hidden="true" > 102 languages </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/Mitochondrium" title="Mitochondrium – Afrikaans" lang="af" hreflang="af" data-title="Mitochondrium" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%85%D9%8A%D8%AA%D9%88%D9%83%D9%86%D8%AF%D8%B1%D9%8A%D9%88%D9%86" title="ميتوكندريون – Arabic" lang="ar" hreflang="ar" data-title="ميتوكندريون" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Aragonese" lang="an" hreflang="an" data-title="Mitocondria" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Mitoxondri" title="Mitoxondri – Azerbaijani" lang="az" hreflang="az" data-title="Mitoxondri" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AE%E0%A6%BE%E0%A6%87%E0%A6%9F%E0%A7%8B%E0%A6%95%E0%A6%A8%E0%A7%8D%E0%A6%A1%E0%A7%8D%E0%A6%B0%E0%A6%BF%E0%A6%AF%E0%A6%BC%E0%A6%BE" title="মাইটোকন্ড্রিয়া – Bangla" lang="bn" hreflang="bn" data-title="মাইটোকন্ড্রিয়া" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/So%C3%A0%E2%81%BF-lia%CC%8Dp-th%C3%A9" title="Soàⁿ-lia̍p-thé – Minnan" lang="nan" hreflang="nan" data-title="Soàⁿ-lia̍p-thé" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9C%D1%96%D1%82%D0%B0%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D1%8B%D1%8F" title="Мітахондрыя – Belarusian" lang="be" hreflang="be" data-title="Мітахондрыя" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F" title="Митохондрия – Bulgarian" lang="bg" hreflang="bg" data-title="Митохондрия" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Mitohondrija" title="Mitohondrija – Bosnian" lang="bs" hreflang="bs" data-title="Mitohondrija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/Mitokondrienn" title="Mitokondrienn – Breton" lang="br" hreflang="br" data-title="Mitokondrienn" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target">Brezhoneg</a></li><li class="interlanguage-link interwiki-ca badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ca.wikipedia.org/wiki/Mitocondri" title="Mitocondri – Catalan" lang="ca" hreflang="ca" data-title="Mitocondri" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://cs.wikipedia.org/wiki/Mitochondrie" title="Mitochondrie – Czech" lang="cs" hreflang="cs" data-title="Mitochondrie" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Welsh" lang="cy" hreflang="cy" data-title="Mitocondria" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-da badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://da.wikipedia.org/wiki/Mitokondrie" title="Mitokondrie – Danish" lang="da" hreflang="da" data-title="Mitokondrie" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Mitochondrium" title="Mitochondrium – German" lang="de" hreflang="de" data-title="Mitochondrium" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Mitokonder" title="Mitokonder – Estonian" lang="et" hreflang="et" data-title="Mitokonder" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9C%CE%B9%CF%84%CE%BF%CF%87%CF%8C%CE%BD%CE%B4%CF%81%CE%B9%CE%BF" title="Μιτοχόνδριο – Greek" lang="el" hreflang="el" data-title="Μιτοχόνδριο" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Spanish" lang="es" hreflang="es" data-title="Mitocondria" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Mitokondrio" title="Mitokondrio – Esperanto" lang="eo" hreflang="eo" data-title="Mitokondrio" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-ext mw-list-item"><a href="https://ext.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Extremaduran" lang="ext" hreflang="ext" data-title="Mitocondria" data-language-autonym="Estremeñu" data-language-local-name="Extremaduran" class="interlanguage-link-target">Estremeñu</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Mitokondrio" title="Mitokondrio – Basque" lang="eu" hreflang="eu" data-title="Mitokondrio" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%85%DB%8C%D8%AA%D9%88%DA%A9%D9%86%D8%AF%D8%B1%DB%8C" title="میتوکندری – Persian" lang="fa" hreflang="fa" data-title="میتوکندری" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Mitochondrie" title="Mitochondrie – French" lang="fr" hreflang="fr" data-title="Mitochondrie" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Mitogondrion" title="Mitogondrion – Western Frisian" lang="fy" hreflang="fy" data-title="Mitogondrion" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target">Frysk</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Miteacoindre" title="Miteacoindre – Irish" lang="ga" hreflang="ga" data-title="Miteacoindre" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gv mw-list-item"><a href="https://gv.wikipedia.org/wiki/Mitochondrion" title="Mitochondrion – Manx" lang="gv" hreflang="gv" data-title="Mitochondrion" data-language-autonym="Gaelg" data-language-local-name="Manx" class="interlanguage-link-target">Gaelg</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Galician" lang="gl" hreflang="gl" data-title="Mitocondria" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/Sien-lia%CC%8Dp-th%C3%AD" title="Sien-lia̍p-thí – Hakka Chinese" lang="hak" hreflang="hak" data-title="Sien-lia̍p-thí" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target">客家語 / Hak-kâ-ngî</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%AF%B8%ED%86%A0%EC%BD%98%EB%93%9C%EB%A6%AC%EC%95%84" title="미토콘드리아 – Korean" lang="ko" hreflang="ko" data-title="미토콘드리아" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%84%D5%AB%D5%BF%D5%B8%D6%84%D5%B8%D5%B6%D5%A4%D6%80%D5%AB%D5%B8%D6%82%D5%B4%D5%B6%D5%A5%D6%80" title="Միտոքոնդրիումներ – Armenian" lang="hy" hreflang="hy" data-title="Միտոքոնդրիումներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B8%E0%A5%82%E0%A4%A4%E0%A5%8D%E0%A4%B0%E0%A4%95%E0%A4%A3%E0%A4%BF%E0%A4%95%E0%A4%BE" title="सूत्रकणिका – Hindi" lang="hi" hreflang="hi" data-title="सूत्रकणिका" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Mitohondrij" title="Mitohondrij – Croatian" lang="hr" hreflang="hr" data-title="Mitohondrij" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Mitokondrio" title="Mitokondrio – Ido" lang="io" hreflang="io" data-title="Mitokondrio" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target">Ido</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Mitokondria" title="Mitokondria – Indonesian" lang="id" hreflang="id" data-title="Mitokondria" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Hvatberi" title="Hvatberi – Icelandic" lang="is" hreflang="is" data-title="Hvatberi" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Mitocondrio" title="Mitocondrio – Italian" lang="it" hreflang="it" data-title="Mitocondrio" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%9E%D7%99%D7%98%D7%95%D7%9B%D7%95%D7%A0%D7%93%D7%A8%D7%99%D7%95%D7%9F" title="מיטוכונדריון – Hebrew" lang="he" hreflang="he" data-title="מיטוכונדריון" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Mitokondria" title="Mitokondria – Javanese" lang="jv" hreflang="jv" data-title="Mitokondria" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%AE%E0%B3%88%E0%B2%9F%E0%B3%8B%E0%B2%95%E0%B2%BE%E0%B2%82%E0%B2%A1%E0%B3%8D%E0%B2%B0%E0%B2%BF%E0%B2%AF%E0%B2%A8%E0%B3%8D" title="ಮೈಟೋಕಾಂಡ್ರಿಯನ್ – Kannada" lang="kn" hreflang="kn" data-title="ಮೈಟೋಕಾಂಡ್ರಿಯನ್" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%9B%E1%83%98%E1%83%A2%E1%83%9D%E1%83%A5%E1%83%9D%E1%83%9C%E1%83%93%E1%83%A0%E1%83%98%E1%83%90" title="მიტოქონდრია – Georgian" lang="ka" hreflang="ka" data-title="მიტოქონდრია" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F" title="Митохондрия – Kazakh" lang="kk" hreflang="kk" data-title="Митохондрия" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Mitokondri" title="Mitokondri – Haitian Creole" lang="ht" hreflang="ht" data-title="Mitokondri" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target">Kreyòl ayisyen</a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/M%C3%AEtokondr%C3%AE" title="Mîtokondrî – Kurdish" lang="ku" hreflang="ku" data-title="Mîtokondrî" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target">Kurdî</a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F%D0%BB%D0%B0%D1%80" title="Митохондриялар – Kyrgyz" lang="ky" hreflang="ky" data-title="Митохондриялар" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target">Кыргызча</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Mitochondrium" title="Mitochondrium – Latin" lang="la" hreflang="la" data-title="Mitochondrium" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Mitohondrijs" title="Mitohondrijs – Latvian" lang="lv" hreflang="lv" data-title="Mitohondrijs" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Mitochondrie" title="Mitochondrie – Luxembourgish" lang="lb" hreflang="lb" data-title="Mitochondrie" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target">Lëtzebuergesch</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Mitochondrija" title="Mitochondrija – Lithuanian" lang="lt" hreflang="lt" data-title="Mitochondrija" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Mitocondrio" title="Mitocondrio – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Mitocondrio" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target">Lingua Franca Nova</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Mitokondrium" title="Mitokondrium – Hungarian" lang="hu" hreflang="hu" data-title="Mitokondrium" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%98%D0%B0" title="Митохондрија – Macedonian" lang="mk" hreflang="mk" data-title="Митохондрија" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%AE%E0%B5%88%E0%B4%B1%E0%B5%8D%E0%B4%B1%E0%B5%8B%E0%B4%95%E0%B5%8B%E0%B5%BA%E0%B4%9F%E0%B5%8D%E0%B4%B0%E0%B4%BF%E0%B4%AF" title="മൈറ്റോകോൺട്രിയ – Malayalam" lang="ml" hreflang="ml" data-title="മൈറ്റോകോൺട്രിയ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%A4%E0%A4%82%E0%A4%A4%E0%A5%82%E0%A4%95%E0%A4%A3%E0%A4%BF%E0%A4%95%E0%A4%BE" title="तंतूकणिका – Marathi" lang="mr" hreflang="mr" data-title="तंतूकणिका" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target">मराठी</a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%9B%E1%83%98%E1%83%A2%E1%83%9D%E1%83%A5%E1%83%9D%E1%83%9C%E1%83%93%E1%83%A0%E1%83%98%E1%83%90" title="მიტოქონდრია – Mingrelian" lang="xmf" hreflang="xmf" data-title="მიტოქონდრია" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target">მარგალური</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Mitokondrion" title="Mitokondrion – Malay" lang="ms" hreflang="ms" data-title="Mitokondrion" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-cdo mw-list-item"><a href="https://cdo.wikipedia.org/wiki/Si%C3%A1ng-l%C4%83k-t%C4%81%CC%A4" title="Siáng-lăk-tā̤ – Mindong" lang="cdo" hreflang="cdo" data-title="Siáng-lăk-tā̤" data-language-autonym="閩東語 / Mìng-dĕ̤ng-ngṳ̄" data-language-local-name="Mindong" class="interlanguage-link-target">閩東語 / Mìng-dĕ̤ng-ngṳ̄</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Mitochondrion" title="Mitochondrion – Dutch" lang="nl" hreflang="nl" data-title="Mitochondrion" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%9F%E3%83%88%E3%82%B3%E3%83%B3%E3%83%89%E3%83%AA%E3%82%A2" title="ミトコンドリア – Japanese" lang="ja" hreflang="ja" data-title="ミトコンドリア" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/Mitochondrium" title="Mitochondrium – Northern Frisian" lang="frr" hreflang="frr" data-title="Mitochondrium" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target">Nordfriisk</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Mitokondrium" title="Mitokondrium – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Mitokondrium" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Mitokondrium" title="Mitokondrium – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Mitokondrium" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Mitocondria" title="Mitocondria – Occitan" lang="oc" hreflang="oc" data-title="Mitocondria" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Mitoxondriya" title="Mitoxondriya – Uzbek" lang="uz" hreflang="uz" data-title="Mitoxondriya" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%A4%E0%A9%B0%E0%A8%A6%E0%A8%95%E0%A9%81%E0%A9%B1%E0%A8%95%E0%A8%B0%E0%A8%BE" title="ਤੰਦਕੁੱਕਰਾ – Punjabi" lang="pa" hreflang="pa" data-title="ਤੰਦਕੁੱਕਰਾ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target">ਪੰਜਾਬੀ</a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D9%85%D8%A7%D8%A6%D9%B9%D9%88%DA%A9%D9%88%D9%86%DA%88%D8%B1%DB%8C%D8%A7" title="مائٹوکونڈریا – Western Punjabi" lang="pnb" hreflang="pnb" data-title="مائٹوکونڈریا" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target">پنجابی</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%85%D8%A7%D9%8A%D8%AA%D9%88%DA%A9%D8%A7%D9%86%D8%AF%D8%B1%D9%8A%D8%A7" title="مايتوکاندريا – Pashto" lang="ps" hreflang="ps" data-title="مايتوکاندريا" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-km mw-list-item"><a href="https://km.wikipedia.org/wiki/%E1%9E%98%E1%9E%B8%E1%9E%8F%E1%9E%BC%E1%9E%80%E1%9E%BB%E1%9E%84%E1%9E%91%E1%9F%92%E1%9E%9A%E1%9E%B8" title="មីតូកុងទ្រី – Khmer" lang="km" hreflang="km" data-title="មីតូកុងទ្រី" data-language-autonym="ភាសាខ្មែរ" data-language-local-name="Khmer" class="interlanguage-link-target">ភាសាខ្មែរ</a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Mitochondrium" title="Mitochondrium – Low German" lang="nds" hreflang="nds" data-title="Mitochondrium" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target">Plattdüütsch</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Mitochondrium" title="Mitochondrium – Polish" lang="pl" hreflang="pl" data-title="Mitochondrium" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Mitoc%C3%B4ndria" title="Mitocôndria – Portuguese" lang="pt" hreflang="pt" data-title="Mitocôndria" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Mitocondrie" title="Mitocondrie – Romanian" lang="ro" hreflang="ro" data-title="Mitocondrie" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-rue mw-list-item"><a href="https://rue.wikipedia.org/wiki/%D0%9C%D1%96%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D1%96%D1%8F" title="Мітохондрія – Rusyn" lang="rue" hreflang="rue" data-title="Мітохондрія" data-language-autonym="Русиньскый" data-language-local-name="Rusyn" class="interlanguage-link-target">Русиньскый</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F" title="Митохондрия – Russian" lang="ru" hreflang="ru" data-title="Митохондрия" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sah mw-list-item"><a href="https://sah.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F" title="Митохондрия – Yakut" lang="sah" hreflang="sah" data-title="Митохондрия" data-language-autonym="Саха тыла" data-language-local-name="Yakut" class="interlanguage-link-target">Саха тыла</a></li><li class="interlanguage-link interwiki-sa mw-list-item"><a href="https://sa.wikipedia.org/wiki/%E0%A4%AE%E0%A4%BE%E0%A4%87%E0%A4%9F%E0%A5%8B%E0%A4%95%E0%A5%89%E0%A4%A8%E0%A5%8D%E0%A4%A1%E0%A5%8D%E0%A4%B0%E0%A4%BF%E0%A4%AF%E0%A4%BE" title="माइटोकॉन्ड्रिया – Sanskrit" lang="sa" hreflang="sa" data-title="माइटोकॉन्ड्रिया" data-language-autonym="संस्कृतम्" data-language-local-name="Sanskrit" class="interlanguage-link-target">संस्कृतम्</a></li><li class="interlanguage-link interwiki-sat mw-list-item"><a href="https://sat.wikipedia.org/wiki/%E1%B1%A2%E1%B1%9F%E1%B1%AD%E1%B1%B4%E1%B1%B3%E1%B1%A0%E1%B1%B3%E1%B1%B1%E1%B1%B0%E1%B1%A8%E1%B1%A4%E1%B1%AD%E1%B1%9F" title="ᱢᱟᱭᱴᱳᱠᱳᱱᱰᱨᱤᱭᱟ – Santali" lang="sat" hreflang="sat" data-title="ᱢᱟᱭᱴᱳᱠᱳᱱᱰᱨᱤᱭᱟ" data-language-autonym="ᱥᱟᱱᱛᱟᱲᱤ" data-language-local-name="Santali" class="interlanguage-link-target">ᱥᱟᱱᱛᱟᱲᱤ</a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Mitochondrion" title="Mitochondrion – Scots" lang="sco" hreflang="sco" data-title="Mitochondrion" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target">Scots</a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Mitokondri" title="Mitokondri – Albanian" lang="sq" hreflang="sq" data-title="Mitokondri" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target">Shqip</a></li><li class="interlanguage-link interwiki-scn mw-list-item"><a href="https://scn.wikipedia.org/wiki/Mitucondri" title="Mitucondri – Sicilian" lang="scn" hreflang="scn" data-title="Mitucondri" data-language-autonym="Sicilianu" data-language-local-name="Sicilian" class="interlanguage-link-target">Sicilianu</a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%B8%E0%B6%BA%E0%B7%92%E0%B6%A7%E0%B7%9C%E0%B6%9A%E0%B7%9C%E0%B6%B1%E0%B7%8A%E0%B6%A9%E0%B7%8A%E2%80%8D%E0%B6%BB%E0%B7%92%E0%B6%BA%E0%B7%8F" title="මයිටොකොන්ඩ්‍රියා – Sinhala" lang="si" hreflang="si" data-title="මයිටොකොන්ඩ්‍රියා" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Mitochondria" title="Mitochondria – Simple English" lang="en-simple" hreflang="en-simple" data-title="Mitochondria" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Mitochondria" title="Mitochondria – Slovak" lang="sk" hreflang="sk" data-title="Mitochondria" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mitohondrij" title="Mitohondrij – Slovenian" lang="sl" hreflang="sl" data-title="Mitohondrij" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%85%D8%A7%DB%8C%D8%AA%DB%86%DA%A9%DB%86%D9%86%D8%AF%D8%B1%DB%8C%D8%A7" title="مایتۆکۆندریا – Central Kurdish" lang="ckb" hreflang="ckb" data-title="مایتۆکۆندریا" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%98%D0%B0" title="Митохондрија – Serbian" lang="sr" hreflang="sr" data-title="Митохондрија" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Mitohondrije" title="Mitohondrije – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Mitohondrije" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Mitokondria" title="Mitokondria – Sundanese" lang="su" hreflang="su" data-title="Mitokondria" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target">Sunda</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Mitokondrio" title="Mitokondrio – Finnish" lang="fi" hreflang="fi" data-title="Mitokondrio" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Mitokondrie" title="Mitokondrie – Swedish" lang="sv" hreflang="sv" data-title="Mitokondrie" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Mitokondriyon" title="Mitokondriyon – Tagalog" lang="tl" hreflang="tl" data-title="Mitokondriyon" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%87%E0%AE%B4%E0%AF%88%E0%AE%AE%E0%AE%A3%E0%AE%BF" title="இழைமணி – Tamil" lang="ta" hreflang="ta" data-title="இழைமணி" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D1%8F" title="Митохондрия – Tatar" lang="tt" hreflang="tt" data-title="Митохондрия" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target">Татарча / tatarça</a></li><li class="interlanguage-link interwiki-te mw-list-item"><a href="https://te.wikipedia.org/wiki/%E0%B0%AE%E0%B1%88%E0%B0%9F%E0%B1%8B%E0%B0%95%E0%B0%BE%E0%B0%82%E0%B0%A1%E0%B1%8D%E0%B0%B0%E0%B0%BF%E0%B0%AF%E0%B0%BE" title="మైటోకాండ్రియా – Telugu" lang="te" hreflang="te" data-title="మైటోకాండ్రియా" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target">తెలుగు</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%84%E0%B8%A1%E0%B9%82%E0%B8%97%E0%B8%84%E0%B8%AD%E0%B8%99%E0%B9%80%E0%B8%94%E0%B8%A3%E0%B8%B5%E0%B8%A2" title="ไมโทคอนเดรีย – Thai" lang="th" hreflang="th" data-title="ไมโทคอนเดรีย" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target">ไทย</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Mitokondri" title="Mitokondri – Turkish" lang="tr" hreflang="tr" data-title="Mitokondri" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9C%D1%96%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D1%96%D1%8F" title="Мітохондрія – Ukrainian" lang="uk" hreflang="uk" data-title="Мітохондрія" data-language-autonym="Українська" data-language-local-name="Ukrainian" 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</div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> <div id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. 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Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></div></div> <div id="mw-indicator-pp-default" class="mw-indicator"><div class="mw-parser-output"><a href="/wiki/Wikipedia:Protection_policy#semi" title="This article is semi-protected due to vandalism"><img alt="Page semi-protected" src="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/20px-Semi-protection-shackle.svg.png" decoding="async" width="20" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/30px-Semi-protection-shackle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/40px-Semi-protection-shackle.svg.png 2x" data-file-width="512" data-file-height="512" /></a></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle">(Redirected from <a href="/w/index.php?title=Mitochondria&redirect=no" class="mw-redirect" title="Mitochondria">Mitochondria</a>)</div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Organelle in eukaryotic cells responsible for respiration</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">"Mitochondria" redirects here. For the song by Kenichi Suzumura, see <a href="/wiki/Mitochondria_(song)" title="Mitochondria (song)">Mitochondria (song)</a>. For the Canadian band, see <a href="/wiki/Mitochondrion_(band)" title="Mitochondrion (band)">Mitochondrion (band)</a>.</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:beige">Mitochondrion</th></tr><tr><td colspan="2" class="infobox-image"><a href="/wiki/File:Animal_mitochondrion_diagram_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Animal_mitochondrion_diagram_en.svg/250px-Animal_mitochondrion_diagram_en.svg.png" decoding="async" width="250" height="171" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Animal_mitochondrion_diagram_en.svg/375px-Animal_mitochondrion_diagram_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Animal_mitochondrion_diagram_en.svg/500px-Animal_mitochondrion_diagram_en.svg.png 2x" data-file-width="582" data-file-height="399" /></a><div class="infobox-caption">Diagram of animal mitochondrion</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Pronunciation</th><td class="infobox-data"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/ˌmaɪtəˈkɒndriən/</a><a href="#cite_note-1">[1]</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Part of</th><td class="infobox-data"><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a></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/Latin" title="Latin">Latin</a></th><td class="infobox-data">organella</td></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=D008928">D008928</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma63835">63835</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_microanatomy" title="Anatomical terms of microanatomy">Anatomical terms of microanatomy</a><div style="text-align: right;">[<a href="https://www.wikidata.org/wiki/Q39572" class="extiw" title="d:Q39572">edit on Wikidata</a>]</div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><table class="infobox" style="width: 300px; padding: 0; border-collapse:collapse; background:transparent;"><tbody><tr><th colspan="2" class="infobox-above" style="background: rgba(0,0,0,0.1); padding: 5px; width: 290px;"><a href="/wiki/Cell_biology" title="Cell biology">Cell biology</a></th></tr><tr><td colspan="2" class="infobox-subheader" style="font-weight:bold; font-size:125%; background-color: #0073CF; color: #FFF;"><a href="/wiki/Animal_cell" class="mw-redirect" title="Animal cell">Animal cell diagram</a></td></tr><tr><td colspan="2" class="infobox-image" style="padding: 0;"><a href="/wiki/File:Animal_Cell.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/300px-Animal_Cell.svg.png" decoding="async" width="300" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/450px-Animal_Cell.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/600px-Animal_Cell.svg.png 2x" data-file-width="1405" data-file-height="811" /></a><div class="infobox-caption" style="text-align:left; padding: 10px; background: rgba(0,0,0,0.02);">Components of a typical animal cell: <ol><li><a href="/wiki/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></li> <li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a> (dots as part of 5)</li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a></li> <li><a href="/wiki/Rough_endoplasmic_reticulum" class="mw-redirect" title="Rough endoplasmic reticulum">Rough endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> (or, Golgi body)</li> <li><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a></li> <li><a href="/wiki/Smooth_endoplasmic_reticulum" class="mw-redirect" title="Smooth endoplasmic reticulum">Smooth endoplasmic reticulum</a></li> <li><a class="mw-selflink selflink">Mitochondrion</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a> (fluid that contains <a href="/wiki/Organelle" title="Organelle">organelles</a>; with which, comprises <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>)</li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Cell_membrane" title="Cell membrane">Cell membrane</a></li></ol></div> </td></tr></tbody></table> A mitochondrion (<abbr title="plural form">pl.</abbr> mitochondria) is an <a href="/wiki/Organelle" title="Organelle">organelle</a> found in the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a> of most <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, such as <a href="/wiki/Animal" title="Animal">animals</a>, <a href="/wiki/Plant" title="Plant">plants</a> and <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>. Mitochondria have a double <a href="/wiki/Lipid_bilayer" title="Lipid bilayer">membrane</a> structure and use <a href="/wiki/Aerobic_respiration" class="mw-redirect" title="Aerobic respiration">aerobic respiration</a> to generate <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP), which is used throughout the cell as a source of <a href="/wiki/Chemical_energy" title="Chemical energy">chemical energy</a>.<a href="#cite_note-2">[2]</a> They were discovered by <a href="/wiki/Albert_von_K%C3%B6lliker" title="Albert von Kölliker">Albert von Kölliker</a> in 1857<a href="#cite_note-Science_in_the_News-2012-3">[3]</a> in the voluntary muscles of insects. Meaning a thread-like granule, the term mitochondrion was coined by <a href="/wiki/Carl_Benda" title="Carl Benda">Carl Benda</a> in 1898. The mitochondrion is popularly nicknamed the "powerhouse of the cell", a phrase popularized by <a href="/wiki/Philip_Siekevitz" title="Philip Siekevitz">Philip Siekevitz</a> in a 1957 <a href="/wiki/Scientific_American" title="Scientific American">Scientific American</a> article of the same name.<a href="#cite_note-Siekevitz-1957-4">[4]</a> Some cells in some <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellular organisms</a> lack mitochondria (for example, mature mammalian <a href="/wiki/Red_blood_cell" title="Red blood cell">red blood cells</a>). The multicellular <a href="/wiki/Animal" title="Animal">animal</a> <a href="/wiki/Henneguya_zschokkei" title="Henneguya zschokkei">Henneguya salminicola</a> is known to have retained mitochondrion-related organelles despite a complete loss of their mitochondrial genome.<a href="#cite_note-Karnkowska-2016-5">[5]</a><a href="#cite_note-Le_Page-2020-6">[6]</a><a href="#cite_note-Yahalomi-2020-7">[7]</a> A large number of <a href="/wiki/Unicellular_organism" title="Unicellular organism">unicellular organisms</a>, such as <a href="/wiki/Microsporidia" title="Microsporidia">microsporidia</a>, <a href="/wiki/Parabasalid" title="Parabasalid">parabasalids</a> and <a href="/wiki/Diplomonad" title="Diplomonad">diplomonads</a>, have reduced or transformed their mitochondria into other structures,<a href="#cite_note-Henze-2003-8">[8]</a> e.g. <a href="/wiki/Hydrogenosome" title="Hydrogenosome">hydrogenosomes</a> and <a href="/wiki/Mitosome" title="Mitosome">mitosomes</a>.<a href="#cite_note-9">[9]</a> The <a href="/wiki/Oxymonad" title="Oxymonad">oxymonads</a> <a href="/wiki/Monocercomonoides" title="Monocercomonoides">Monocercomonoides</a>, <a href="/wiki/Streblomastix" title="Streblomastix">Streblomastix</a>, and <a href="/w/index.php?title=Blattamonas&action=edit&redlink=1" class="new" title="Blattamonas (page does not exist)">Blattamonas</a> have completely lost their mitochondria.<a href="#cite_note-Karnkowska-2016-5">[5]</a><a href="#cite_note-Novák-2023-10">[10]</a> Mitochondria are commonly between 0.75 and 3 <a href="/wiki/Micrometre" title="Micrometre">μm</a>2 in cross section,<a href="#cite_note-Wiemerslage-2016-11">[11]</a> but vary considerably in size and structure. Unless specifically <a href="/wiki/Staining" title="Staining">stained</a>, they are not visible. In addition to supplying cellular energy, mitochondria are involved in other tasks, such as <a href="/wiki/Cell_signaling" title="Cell signaling">signaling</a>, <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">cellular differentiation</a>, and <a href="/wiki/Apoptosis" title="Apoptosis">cell death</a>, as well as maintaining control of the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> and <a href="/wiki/Cell_growth" title="Cell growth">cell growth</a>.<a href="#cite_note-McBride-2006-12">[12]</a> <a href="/wiki/Mitochondrial_biogenesis" title="Mitochondrial biogenesis">Mitochondrial biogenesis</a> is in turn temporally coordinated with these cellular processes.<a href="#cite_note-Valero-2014-13">[13]</a><a href="#cite_note-Sanchis-Gomar-2014-14">[14]</a> Mitochondria have been implicated in several human disorders and conditions, such as <a href="/wiki/Mitochondrial_disease" title="Mitochondrial disease">mitochondrial diseases</a>,<a href="#cite_note-15">[15]</a> <a href="/wiki/Heart_failure" title="Heart failure">cardiac dysfunction</a>,<a href="#cite_note-16">[16]</a> heart failure<a href="#cite_note-Dorn-2015-17">[17]</a> and <a href="/wiki/Autism" title="Autism">autism</a>.<a href="#cite_note-Griffiths-2017-18">[18]</a> The number of mitochondria in a cell can vary widely by <a href="/wiki/Organism" title="Organism">organism</a>, <a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissue</a>, and cell type. A mature red blood cell has no mitochondria,<a href="#cite_note-Ney-2011-19">[19]</a> whereas a <a href="/wiki/Hepatocyte" title="Hepatocyte">liver cell</a> can have more than 2000.<a href="#cite_note-Alberts-2005-20">[20]</a><a href="#cite_note-Voet-2006-21">[21]</a> The mitochondrion is composed of compartments that carry out specialized functions. These compartments or regions include the outer membrane, <a href="/wiki/Intermembrane_space" title="Intermembrane space">intermembrane space</a>, <a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">inner membrane</a>, <a href="/wiki/Crista" title="Crista">cristae</a>, and <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">matrix</a>. Although most of a eukaryotic cell's <a href="/wiki/DNA" title="DNA">DNA</a> is contained in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>, the mitochondrion has <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">its own genome</a> ("mitogenome") that is substantially similar to <a href="/wiki/Bacteria" title="Bacteria">bacterial</a> genomes.<a href="#cite_note-22">[22]</a> This finding has led to general acceptance of the <a href="/wiki/Symbiogenesis" title="Symbiogenesis">endosymbiotic hypothesis</a> - that free-living prokaryotic ancestors of modern mitochondria permanently fused with eukaryotic cells in the distant past, evolving such that modern animals, plants, fungi, and other eukaryotes are able to <a href="/wiki/Cellular_respiration" title="Cellular respiration">respire</a> to generate <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">cellular energy</a>.<a href="#cite_note-Gabaldón-2021-23">[23]</a> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2></div> Mitochondria may have a number of different shapes.<a href="#cite_note-British_Society_for_Cell_Biology-24">[24]</a> A mitochondrion contains outer and inner membranes composed of <a href="/wiki/Phospholipid_bilayer" class="mw-redirect" title="Phospholipid bilayer">phospholipid bilayers</a> and <a href="/wiki/Protein" title="Protein">proteins</a>.<a href="#cite_note-Alberts-2005-20">[20]</a> The two membranes have different properties. Because of this double-membraned organization, there are five distinct parts to a mitochondrion: <ol><li>The outer mitochondrial membrane,</li> <li>The intermembrane space (the space between the outer and inner membranes),</li> <li>The inner mitochondrial membrane,</li> <li>The <a href="/wiki/Crista" title="Crista">cristae</a> space (formed by infoldings of the inner membrane), and</li> <li>The <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">matrix</a> (space within the inner membrane), which is a fluid.</li></ol> Mitochondria have folding to increase surface area, which in turn increases ATP (adenosine triphosphate) production. Mitochondria stripped of their outer membrane are called <a href="/wiki/Mitoplast" title="Mitoplast">mitoplasts</a>. <div class="mw-heading mw-heading3"><h3 id="Outer_membrane">Outer membrane</h3></div> The outer mitochondrial membrane, which encloses the entire organelle, is 60 to 75 <a href="/wiki/Angstrom" title="Angstrom">angstroms</a> (Å) thick. It has a protein-to-phospholipid ratio similar to that of the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> (about 1:1 by weight). It contains large numbers of <a href="/wiki/Integral_membrane_protein" title="Integral membrane protein">integral membrane proteins</a> called <a href="/wiki/Porin_(protein)" title="Porin (protein)">porins</a>. A major trafficking protein is the pore-forming <a href="/wiki/Voltage-dependent_anion_channel" title="Voltage-dependent anion channel">voltage-dependent anion channel</a> (VDAC). The <a href="/wiki/Voltage-dependent_anion_channel#Biological_function" title="Voltage-dependent anion channel">VDAC</a> is the primary transporter of <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a>, <a href="/wiki/Ion" title="Ion">ions</a> and <a href="/wiki/Metabolite" title="Metabolite">metabolites</a> between the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> and the intermembrane space.<a href="#cite_note-Blachly-Dyson-2001-25">[25]</a><a href="#cite_note-Hoogenboom-2007-26">[26]</a> It is formed as a <a href="/wiki/Beta_barrel" title="Beta barrel">beta barrel</a> that spans the outer membrane, similar to that in the <a href="/wiki/Gram-negative" class="mw-redirect" title="Gram-negative">gram-negative</a> <a href="/wiki/Bacterial_outer_membrane" title="Bacterial outer membrane">bacterial outer membrane</a>.<a href="#cite_note-Zeth-2010-27">[27]</a> Larger proteins can enter the mitochondrion if a signaling sequence at their <a href="/wiki/N-terminus" title="N-terminus">N-terminus</a> binds to a large multisubunit <a href="/wiki/Mitochondrial_membrane_transport_protein" title="Mitochondrial membrane transport protein">protein</a> called <a href="/wiki/Translocase_of_the_outer_membrane" title="Translocase of the outer membrane">translocase in the outer membrane</a>, which then <a href="/wiki/Active_transport" title="Active transport">actively moves</a> them across the membrane.<a href="#cite_note-Herrmann-2000-28">[28]</a> Mitochondrial <a href="/wiki/Protein_precursor" title="Protein precursor">pro-proteins</a> are imported through specialised translocation complexes. The outer membrane also contains <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> involved in such diverse activities as the elongation of <a href="/wiki/Fatty_acid" title="Fatty acid">fatty acids</a>, <a href="/wiki/Oxidation" class="mw-redirect" title="Oxidation">oxidation</a> of <a href="/wiki/Epinephrine" class="mw-redirect" title="Epinephrine">epinephrine</a>, and the <a href="/wiki/Biodegradation" title="Biodegradation">degradation</a> of <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a>. These enzymes include <a href="/wiki/Monoamine_oxidase" title="Monoamine oxidase">monoamine oxidase</a>, <a href="/wiki/Rotenone" title="Rotenone">rotenone</a>-insensitive NADH-cytochrome c-reductase, <a href="/wiki/Kynurenine" title="Kynurenine">kynurenine</a> <a href="/wiki/Hydroxylase" class="mw-redirect" title="Hydroxylase">hydroxylase</a> and fatty acid Co-A <a href="/wiki/Ligase" title="Ligase">ligase</a>. Disruption of the outer membrane permits proteins in the intermembrane space to leak into the cytosol, leading to cell death.<a href="#cite_note-Chipuk-2006-29">[29]</a> The outer mitochondrial membrane can associate with the endoplasmic reticulum (ER) membrane, in a structure called MAM (mitochondria-associated ER-membrane). This is important in the ER-mitochondria calcium signaling and is involved in the transfer of lipids between the ER and mitochondria.<a href="#cite_note-Hayashi-2009-30">[30]</a> Outside the outer membrane are small (diameter: 60 Å) particles named sub-units of Parson. <div class="mw-heading mw-heading3"><h3 id="Intermembrane_space">Intermembrane space</h3></div> The mitochondrial intermembrane space is the space between the outer membrane and the inner membrane. It is also known as perimitochondrial space. Because the outer membrane is freely permeable to small molecules, the concentrations of small molecules, such as ions and sugars, in the intermembrane space is the same as in the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>.<a href="#cite_note-Alberts-2005-20">[20]</a> However, large proteins must have a specific signaling sequence to be transported across the outer membrane, so the protein composition of this space is different from the protein composition of the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>. One <a href="/wiki/Protein" title="Protein">protein</a> that is localized to the intermembrane space in this way is <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome c</a>.<a href="#cite_note-Chipuk-2006-29">[29]</a> <div class="mw-heading mw-heading3"><h3 id="Inner_membrane">Inner membrane</h3></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/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">Inner mitochondrial membrane</a></div> The inner mitochondrial membrane contains proteins with three types of functions:<a href="#cite_note-Alberts-2005-20">[20]</a> <ol><li>Those that perform the <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport chain</a> <a href="/wiki/Redox" title="Redox">redox</a> reactions</li> <li><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a>, which generates <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> in the matrix</li> <li>Specific <a href="/wiki/Membrane_transport_protein" title="Membrane transport protein">transport proteins</a> that regulate <a href="/wiki/Metabolite" title="Metabolite">metabolite</a> passage into and out of the <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">mitochondrial matrix</a></li></ol> It contains more than 151 different <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptides</a>, and has a very high protein-to-phospholipid ratio (more than 3:1 by weight, which is about 1 protein for 15 phospholipids). The inner membrane is home to around 1/5 of the total protein in a mitochondrion.<a href="#cite_note-Schenkel-2014-31">[31]</a> Additionally, the inner membrane is rich in an unusual phospholipid, <a href="/wiki/Cardiolipin" title="Cardiolipin">cardiolipin</a>. This phospholipid was originally discovered in <a href="/wiki/Bos_taurus" class="mw-redirect" title="Bos taurus">cow</a> hearts in 1942, and is usually characteristic of mitochondrial and bacterial plasma membranes.<a href="#cite_note-McMillin-2002-32">[32]</a> Cardiolipin contains four fatty acids rather than two, and may help to make the inner membrane impermeable,<a href="#cite_note-Alberts-2005-20">[20]</a> and its disruption can lead to multiple clinical disorders including neurological disorders and cancer.<a href="#cite_note-33">[33]</a> Unlike the outer membrane, the inner membrane does not contain porins, and is highly impermeable to all molecules. Almost all ions and molecules require special membrane transporters to enter or exit the matrix. Proteins are ferried into the matrix via the <a href="/wiki/Translocase_of_the_inner_membrane" title="Translocase of the inner membrane">translocase of the inner membrane</a> (TIM) complex or via <a href="/wiki/OXA1L" title="OXA1L">OXA1L</a>.<a href="#cite_note-Herrmann-2000-28">[28]</a> In addition, there is a membrane potential across the inner membrane, formed by the action of the <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> of the <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport chain</a>. Inner membrane <a href="/wiki/Mitochondrial_fusion" title="Mitochondrial fusion">fusion</a> is mediated by the inner membrane protein <a href="/wiki/OPA1" class="mw-redirect" title="OPA1">OPA1</a>.<a href="#cite_note-Youle-2012-34">[34]</a> <div class="mw-heading mw-heading4"><h4 id="Cristae">Cristae</h4></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:MitochondrionCAM.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/MitochondrionCAM.jpg/250px-MitochondrionCAM.jpg" decoding="async" width="250" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/MitochondrionCAM.jpg/375px-MitochondrionCAM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/MitochondrionCAM.jpg/500px-MitochondrionCAM.jpg 2x" data-file-width="618" data-file-height="409" /></a><figcaption>Cross-sectional image of cristae in a <a href="/wiki/Rat" title="Rat">rat</a> liver mitochondrion to demonstrate the likely 3D structure and relationship to the inner membrane</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Crista" title="Crista">Crista</a></div> The inner mitochondrial membrane is compartmentalized into numerous folds called <a href="/wiki/Crista" title="Crista">cristae</a>, which expand the surface area of the inner mitochondrial membrane, enhancing its ability to produce ATP. For typical liver mitochondria, the area of the inner membrane is about five times as large as that of the outer membrane. This ratio is variable and mitochondria from cells that have a greater demand for ATP, such as muscle cells, contain even more cristae. Mitochondria within the same cell can have substantially different crista-density, with the ones that are required to produce more energy having much more crista-membrane surface.<a href="#cite_note-Cserép-2018-35">[35]</a> These folds are studded with small round bodies known as <a href="/wiki/F-ATPase" title="F-ATPase">F1 particles</a> or oxysomes.<a href="#cite_note-Mannella-2006-36">[36]</a> <div class="mw-heading mw-heading3"><h3 id="Matrix">Matrix</h3></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/Mitochondrial_matrix" title="Mitochondrial matrix">Mitochondrial matrix</a></div> The matrix is the space enclosed by the inner membrane. It contains about 2/3 of the total proteins in a mitochondrion.<a href="#cite_note-Alberts-2005-20">[20]</a> The matrix is important in the production of ATP with the aid of the ATP synthase contained in the inner membrane. The matrix contains a highly concentrated mixture of hundreds of enzymes, special mitochondrial <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a>, <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a>, and several copies of the <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> <a href="/wiki/Genome" title="Genome">genome</a>. Of the enzymes, the major functions include oxidation of <a href="/wiki/Pyruvate" class="mw-redirect" title="Pyruvate">pyruvate</a> and <a href="/wiki/Fatty_acids" class="mw-redirect" title="Fatty acids">fatty acids</a>, and the <a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</a>.<a href="#cite_note-Alberts-2005-20">[20]</a> The DNA molecules are packaged into nucleoids by proteins, one of which is <a href="/wiki/TFAM" title="TFAM">TFAM</a>.<a href="#cite_note-37">[37]</a> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2></div> The most prominent roles of mitochondria are to produce the energy currency of the cell, <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> (i.e., phosphorylation of <a href="/wiki/Adenosine_diphosphate" title="Adenosine diphosphate">ADP</a>), through respiration and to regulate cellular <a href="/wiki/Metabolism" title="Metabolism">metabolism</a>.<a href="#cite_note-Voet-2006-21">[21]</a> The central set of reactions involved in ATP production are collectively known as the <a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</a>, or the <a href="/wiki/Hans_Adolf_Krebs" class="mw-redirect" title="Hans Adolf Krebs">Krebs</a> cycle, and <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a>. However, the mitochondrion has many other functions in addition to the production of ATP. <div class="mw-heading mw-heading3"><h3 id="Energy_conversion">Energy conversion</h3></div> A dominant role for the mitochondria is the production of ATP, as reflected by the large number of proteins in the inner membrane for this task. This is done by oxidizing the major products of <a href="/wiki/Glucose" title="Glucose">glucose</a>: <a href="/wiki/Pyruvate" class="mw-redirect" title="Pyruvate">pyruvate</a>, and <a href="/wiki/NADH" class="mw-redirect" title="NADH">NADH</a>, which are produced in the cytosol.<a href="#cite_note-Voet-2006-21">[21]</a> This type of <a href="/wiki/Cellular_respiration" title="Cellular respiration">cellular respiration</a>, known as <a href="/wiki/Aerobic_respiration" class="mw-redirect" title="Aerobic respiration">aerobic respiration</a>, is dependent on the presence of <a href="/wiki/Oxygen" title="Oxygen">oxygen</a>. When oxygen is limited, the glycolytic products will be metabolized by <a href="/wiki/Fermentation_(biochemistry)" class="mw-redirect" title="Fermentation (biochemistry)">anaerobic fermentation</a>, a process that is independent of the mitochondria.<a href="#cite_note-Voet-2006-21">[21]</a> The production of ATP from glucose and oxygen has an approximately 13-times higher yield during aerobic respiration compared to fermentation.<a href="#cite_note-38">[38]</a> Plant mitochondria can also produce a limited amount of ATP either by breaking the sugar produced during photosynthesis or without oxygen by using the alternate substrate <a href="/wiki/Nitrite" title="Nitrite">nitrite</a>.<a href="#cite_note-Stoimenova-2007-39">[39]</a> ATP crosses out through the inner membrane with the help of a <a href="/wiki/ATP%E2%80%93ADP_translocase" class="mw-redirect" title="ATP–ADP translocase">specific protein</a>, and across the outer membrane via <a href="/wiki/Porin_(protein)" title="Porin (protein)">porins</a>.<a href="#cite_note-Neupert-1997-40">[40]</a> After conversion of ATP to <a href="/wiki/Adenosine_diphosphate" title="Adenosine diphosphate">ADP</a> by <a href="/wiki/Dephosphorylation" title="Dephosphorylation">dephosphorylation</a> that releases energy, ADP returns via the same route. <div class="mw-heading mw-heading4"><h4 id="Pyruvate_and_the_citric_acid_cycle">Pyruvate and the citric acid cycle</h4></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/Citric_acid_cycle" title="Citric acid cycle">Citric acid cycle</a></div> <a href="/wiki/Pyruvate" class="mw-redirect" title="Pyruvate">Pyruvate</a> molecules produced by <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a> are <a href="/wiki/Active_transport" title="Active transport">actively transported</a> across the inner mitochondrial membrane, and into the matrix where they can either be <a href="/wiki/Redox" title="Redox">oxidized</a> and combined with <a href="/wiki/Coenzyme_A" title="Coenzyme A">coenzyme A</a> to form CO2, <a href="/wiki/Acetyl-CoA" title="Acetyl-CoA">acetyl-CoA</a>, and <a href="/wiki/NADH" class="mw-redirect" title="NADH">NADH</a>,<a href="#cite_note-Voet-2006-21">[21]</a> or they can be <a href="/wiki/Carboxylated" class="mw-redirect" title="Carboxylated">carboxylated</a> (by <a href="/wiki/Pyruvate_carboxylase" title="Pyruvate carboxylase">pyruvate carboxylase</a>) to form oxaloacetate. This latter reaction "fills up" the amount of oxaloacetate in the citric acid cycle and is therefore an <a href="/wiki/Anaplerotic_reaction" class="mw-redirect" title="Anaplerotic reaction">anaplerotic reaction</a>, increasing the cycle's capacity to metabolize acetyl-CoA when the tissue's energy needs (e.g., in <a href="/wiki/Striated_muscle_tissue" title="Striated muscle tissue">muscle</a>) are suddenly increased by activity.<a href="#cite_note-Stryer-1995-41">[41]</a> In the citric acid cycle, all the intermediates (e.g. <a href="/wiki/Citrate" class="mw-redirect" title="Citrate">citrate</a>, <a href="/wiki/Isocitric_acid" title="Isocitric acid">iso-citrate</a>, <a href="/wiki/Alpha-Ketoglutaric_acid" class="mw-redirect" title="Alpha-Ketoglutaric acid">alpha-ketoglutarate</a>, succinate, <a href="/wiki/Fumaric_acid" title="Fumaric acid">fumarate</a>, <a href="/wiki/Malic_acid" title="Malic acid">malate</a> and oxaloacetate) are regenerated during each turn of the cycle. Adding more of any of these intermediates to the mitochondrion therefore means that the additional amount is retained within the cycle, increasing all the other intermediates as one is converted into the other. Hence, the addition of any one of them to the cycle has an <a href="/wiki/Anaplerotic_reactions" title="Anaplerotic reactions">anaplerotic</a> effect, and its removal has a cataplerotic effect. These anaplerotic and <a href="/w/index.php?title=Cataplerotic&action=edit&redlink=1" class="new" title="Cataplerotic (page does not exist)">cataplerotic</a> reactions will, during the course of the cycle, increase or decrease the amount of oxaloacetate available to combine with acetyl-CoA to form citric acid. This in turn increases or decreases the rate of <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> production by the mitochondrion, and thus the availability of ATP to the cell.<a href="#cite_note-Stryer-1995-41">[41]</a> Acetyl-CoA, on the other hand, derived from pyruvate oxidation, or from the <a href="/wiki/Beta-oxidation" class="mw-redirect" title="Beta-oxidation">beta-oxidation</a> of <a href="/wiki/Fatty_acids" class="mw-redirect" title="Fatty acids">fatty acids</a>, is the only fuel to enter the citric acid cycle. With each turn of the cycle one molecule of acetyl-CoA is consumed for every molecule of oxaloacetate present in the mitochondrial matrix, and is never regenerated. It is the oxidation of the acetate portion of acetyl-CoA that produces CO2 and water, with the energy thus released captured in the form of ATP.<a href="#cite_note-Stryer-1995-41">[41]</a> In the liver, the <a href="/wiki/Carboxylation" title="Carboxylation">carboxylation</a> of <a href="/wiki/Cytosol" title="Cytosol">cytosolic</a> pyruvate into intra-mitochondrial oxaloacetate is an early step in the <a href="/wiki/Gluconeogenesis" title="Gluconeogenesis">gluconeogenic</a> pathway, which converts <a href="/wiki/Lactic_acid" title="Lactic acid">lactate</a> and de-aminated <a href="/wiki/Alanine" title="Alanine">alanine</a> into glucose,<a href="#cite_note-Voet-2006-21">[21]</a><a href="#cite_note-Stryer-1995-41">[41]</a> under the influence of high levels of <a href="/wiki/Glucagon" title="Glucagon">glucagon</a> and/or <a href="/wiki/Epinephrine" class="mw-redirect" title="Epinephrine">epinephrine</a> in the blood.<a href="#cite_note-Stryer-1995-41">[41]</a> Here, the addition of oxaloacetate to the mitochondrion does not have a net anaplerotic effect, as another citric acid cycle intermediate (malate) is immediately removed from the mitochondrion to be converted to cytosolic oxaloacetate, and ultimately to glucose, in a process that is almost the reverse of <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a>.<a href="#cite_note-Stryer-1995-41">[41]</a> The enzymes of the citric acid cycle are located in the mitochondrial matrix, with the exception of <a href="/wiki/Succinate_dehydrogenase" title="Succinate dehydrogenase">succinate dehydrogenase</a>, which is bound to the inner mitochondrial membrane as part of Complex II.<a href="#cite_note-42">[42]</a> The citric acid cycle oxidizes the acetyl-CoA to carbon dioxide, and, in the process, produces reduced cofactors (three molecules of <a href="/wiki/NADH" class="mw-redirect" title="NADH">NADH</a> and one molecule of <a href="/wiki/FADH2" class="mw-redirect" title="FADH2">FADH2</a>) that are a source of electrons for the <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport chain</a>, and a molecule of <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> (which is readily converted to an ATP).<a href="#cite_note-Voet-2006-21">[21]</a> <div class="mw-heading mw-heading4"><h4 id="O2_and_NADH:_energy-releasing_reactions">O2 and NADH: energy-releasing reactions</h4></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">Oxidative phosphorylation</a> and <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport chain</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Electron_transport_chain.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Electron_transport_chain.svg/400px-Electron_transport_chain.svg.png" decoding="async" width="400" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Electron_transport_chain.svg/600px-Electron_transport_chain.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Electron_transport_chain.svg/800px-Electron_transport_chain.svg.png 2x" data-file-width="822" data-file-height="435" /></a><figcaption>Electron transport chain in the mitochondrial intermembrane space</figcaption></figure> The electrons from NADH and FADH2 are transferred to oxygen (O2) and hydrogen (protons) in several steps via an electron transport chain. NADH and FADH2 molecules are produced within the matrix via the citric acid cycle and in the cytoplasm by <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a>. <a href="/wiki/Reducing_equivalent" class="mw-redirect" title="Reducing equivalent">Reducing equivalents</a> from the cytoplasm can be imported via the <a href="/wiki/Malate-aspartate_shuttle" class="mw-redirect" title="Malate-aspartate shuttle">malate-aspartate shuttle</a> system of <a href="/wiki/Antiporter" title="Antiporter">antiporter</a> proteins or fed into the electron transport chain using a <a href="/wiki/Glycerol_phosphate_shuttle" title="Glycerol phosphate shuttle">glycerol phosphate shuttle</a>.<a href="#cite_note-Voet-2006-21">[21]</a> The major energy-releasing reactions<a href="#cite_note-Voet-2004-43">[43]</a><a href="#cite_note-Atkins-2006-44">[44]</a> that make the mitochondrion the "powerhouse of the cell" occur at <a href="/wiki/Electron_transport_chain#Mitochondrial_redox_carriers" title="Electron transport chain">protein complexes I, III and IV</a> in the inner mitochondrial membrane (<a href="/wiki/NADH_dehydrogenase_(ubiquinone)" class="mw-redirect" title="NADH dehydrogenase (ubiquinone)">NADH dehydrogenase (ubiquinone)</a>, <a href="/wiki/Coenzyme_Q_-_cytochrome_c_reductase" class="mw-redirect" title="Coenzyme Q - cytochrome c reductase">cytochrome c reductase</a>, and <a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">cytochrome c oxidase</a>). At <a href="/wiki/Complex_IV" class="mw-redirect" title="Complex IV">complex IV</a>, O2 reacts with the reduced form of iron in <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome c</a>: <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\ce {O2{}+4H+(aq){}+4Fe^{2+}(cyt\,c)->2H2O{}+4Fe^{3+}(cyt\,c)}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <msubsup> <mtext>O</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mspace width="0pt" height="0pt" depth=".2em" /> </mrow> </msubsup> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mn>4</mn> <mspace width="thinmathspace" /> <msup> <mtext>H</mtext> <mrow class="MJX-TeXAtom-ORD"> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>aq</mtext> <mo stretchy="false">)</mo> </mrow> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mn>4</mn> <mspace width="thinmathspace" /> <msup> <mtext>Fe</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mspace width="thinmathspace" /> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>cyt</mtext> <mspace width="thinmathspace" /> <mtext>c</mtext> <mo stretchy="false">)</mo> </mrow> <mo stretchy="false">⟶</mo> <mn>2</mn> <mspace width="thinmathspace" /> <msubsup> <mtext>H</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mspace width="0pt" height="0pt" depth=".2em" /> </mrow> </msubsup> <mtext>O</mtext> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mn>4</mn> <mspace width="thinmathspace" /> <msup> <mtext>Fe</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>3</mn> <mspace width="thinmathspace" /> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>cyt</mtext> <mspace width="thinmathspace" /> <mtext>c</mtext> <mo stretchy="false">)</mo> </mrow> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\ce {O2{}+4H+(aq){}+4Fe^{2+}(cyt\,c)->2H2O{}+4Fe^{3+}(cyt\,c)}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/aaaa54ad146de2576c6f478dad4c20fa622c6842" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:58.063ex; height:3.343ex;" alt="{\displaystyle {\ce {O2{}+4H+(aq){}+4Fe^{2+}(cyt\,c)->2H2O{}+4Fe^{3+}(cyt\,c)}}}"> <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta _{r}G^{o'}=-218{\text{ kJ/mol}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi mathvariant="normal">Δ</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <msup> <mi>G</mi> <mrow class="MJX-TeXAtom-ORD"> <msup> <mi>o</mi> <mo>′</mo> </msup> </mrow> </msup> <mo>=</mo> <mo>−</mo> <mn>218</mn> <mrow class="MJX-TeXAtom-ORD"> <mtext> kJ/mol</mtext> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta _{r}G^{o'}=-218{\text{ kJ/mol}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/58626e5bfd5c127532e3c8151aba4974842f2b43" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:22.602ex; height:3.343ex;" alt="{\displaystyle \Delta _{r}G^{o'}=-218{\text{ kJ/mol}}}"> releasing a lot of <a href="/wiki/Gibbs_free_energy" title="Gibbs free energy">free energy</a><a href="#cite_note-Atkins-2006-44">[44]</a><a href="#cite_note-Voet-2004-43">[43]</a> from the reactants without breaking bonds of an organic fuel. The free energy put in to remove an electron from Fe2+ is released at <a href="/wiki/Complex_III" class="mw-redirect" title="Complex III">complex III</a> when Fe3+ of cytochrome c reacts to oxidize <a href="/wiki/Ubiquinol" title="Ubiquinol">ubiquinol</a> (QH2): <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\ce {2Fe^{3+}(cyt\,c){}+QH2->2Fe^{2+}(cyt\,c){}+Q{}+2H+(aq)}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mspace width="thinmathspace" /> <msup> <mtext>Fe</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>3</mn> <mspace width="thinmathspace" /> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>cyt</mtext> <mspace width="thinmathspace" /> <mtext>c</mtext> <mo stretchy="false">)</mo> </mrow> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <msubsup> <mtext>QH</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mspace width="0pt" height="0pt" depth=".2em" /> </mrow> </msubsup> <mo stretchy="false">⟶</mo> <mn>2</mn> <mspace width="thinmathspace" /> <msup> <mtext>Fe</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mspace width="thinmathspace" /> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>cyt</mtext> <mspace width="thinmathspace" /> <mtext>c</mtext> <mo stretchy="false">)</mo> </mrow> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mtext>Q</mtext> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mn>2</mn> <mspace width="thinmathspace" /> <msup> <mtext>H</mtext> <mrow class="MJX-TeXAtom-ORD"> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>aq</mtext> <mo stretchy="false">)</mo> </mrow> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\ce {2Fe^{3+}(cyt\,c){}+QH2->2Fe^{2+}(cyt\,c){}+Q{}+2H+(aq)}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/94b741f851cbe8a1ce9ee62dcd03bdfd9d870367" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:55.459ex; height:3.343ex;" alt="{\displaystyle {\ce {2Fe^{3+}(cyt\,c){}+QH2->2Fe^{2+}(cyt\,c){}+Q{}+2H+(aq)}}}"> <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta _{r}G^{o'}=-30{\text{ kJ/mol}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi mathvariant="normal">Δ</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <msup> <mi>G</mi> <mrow class="MJX-TeXAtom-ORD"> <msup> <mi>o</mi> <mo>′</mo> </msup> </mrow> </msup> <mo>=</mo> <mo>−</mo> <mn>30</mn> <mrow class="MJX-TeXAtom-ORD"> <mtext> kJ/mol</mtext> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta _{r}G^{o'}=-30{\text{ kJ/mol}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/b79d3440884b3c1eb8b1fd595152d48b06e4a72c" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:21.44ex; height:3.343ex;" alt="{\displaystyle \Delta _{r}G^{o'}=-30{\text{ kJ/mol}}}"> The <a href="/wiki/Ubiquinone" class="mw-redirect" title="Ubiquinone">ubiquinone</a> (Q) generated reacts, in <a href="/wiki/Complex_I" class="mw-redirect" title="Complex I">complex I</a>, with NADH: <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\ce {Q + H+(aq){}+ NADH -> QH2 + NAD+ {}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mtext>Q</mtext> <mo>+</mo> <msup> <mtext>H</mtext> <mrow class="MJX-TeXAtom-ORD"> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> <mo stretchy="false">(</mo> <mtext>aq</mtext> <mo stretchy="false">)</mo> </mrow> <mrow class="MJX-TeXAtom-ORD"> </mrow> <mo>+</mo> <mtext>NADH</mtext> <mo stretchy="false">⟶</mo> <msubsup> <mtext>QH</mtext> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mspace width="0pt" height="0pt" depth=".2em" /> </mrow> </msubsup> <mo>+</mo> <msup> <mtext>NAD</mtext> <mrow class="MJX-TeXAtom-ORD"> <mo>+</mo> </mrow> </msup> <mrow class="MJX-TeXAtom-ORD"> </mrow> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\ce {Q + H+(aq){}+ NADH -> QH2 + NAD+ {}}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/96ddc7445b7e06f4a9ee73a318e88ffd799a22f9" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:41.261ex; height:3.176ex;" alt="{\displaystyle {\ce {Q + H+(aq){}+ NADH -> QH2 + NAD+ {}}}}"> <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta _{r}G^{o'}=-81{\text{ kJ/mol}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi mathvariant="normal">Δ</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <msup> <mi>G</mi> <mrow class="MJX-TeXAtom-ORD"> <msup> <mi>o</mi> <mo>′</mo> </msup> </mrow> </msup> <mo>=</mo> <mo>−</mo> <mn>81</mn> <mrow class="MJX-TeXAtom-ORD"> <mtext> kJ/mol</mtext> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta _{r}G^{o'}=-81{\text{ kJ/mol}}}</annotation> </semantics> </math><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/e296bcbb580c2f1e4936acde7e67b0f32801e855" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:21.44ex; height:3.343ex;" alt="{\displaystyle \Delta _{r}G^{o'}=-81{\text{ kJ/mol}}}"> While the reactions are controlled by an electron transport chain, free electrons are not amongst the reactants or products in the three reactions shown and therefore do not affect the free energy released, which is used to pump <a href="/wiki/Hydrogen_ion" title="Hydrogen ion">protons</a> (H+) into the intermembrane space. This process is efficient, but a small percentage of electrons may prematurely reduce oxygen, forming <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> such as <a href="/wiki/Superoxide" title="Superoxide">superoxide</a>.<a href="#cite_note-Voet-2006-21">[21]</a> This can cause <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a> in the mitochondria and may contribute to the decline in mitochondrial function associated with aging.<a href="#cite_note-Huang-2004-45">[45]</a> As the proton concentration increases in the intermembrane space, a strong <a href="/wiki/Electrochemical_gradient" title="Electrochemical gradient">electrochemical gradient</a> is established across the inner membrane. The protons can return to the matrix through the <a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a> complex, and their potential energy is used to synthesize <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> from ADP and inorganic phosphate (Pi).<a href="#cite_note-Voet-2006-21">[21]</a> This process is called <a href="/wiki/Chemiosmosis" title="Chemiosmosis">chemiosmosis</a>, and was first described by <a href="/wiki/Peter_D._Mitchell" title="Peter D. Mitchell">Peter Mitchell</a>,<a href="#cite_note-Mitchell-1967a-46">[46]</a><a href="#cite_note-Mitchell-1967b-47">[47]</a> who was awarded the 1978 <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a> for his work. Later, part of the 1997 Nobel Prize in Chemistry was awarded to <a href="/wiki/Paul_D._Boyer" title="Paul D. Boyer">Paul D. Boyer</a> and <a href="/wiki/John_E._Walker" title="John E. Walker">John E. Walker</a> for their clarification of the working mechanism of ATP synthase.<a href="#cite_note-48">[48]</a> <div class="mw-heading mw-heading4"><h4 id="Heat_production">Heat production</h4></div> Under certain conditions, protons can re-enter the mitochondrial matrix without contributing to ATP synthesis. This process is known as proton leak or <a href="/wiki/Mitochondrial_uncoupling" class="mw-redirect" title="Mitochondrial uncoupling">mitochondrial uncoupling</a> and is due to the <a href="/wiki/Facilitated_diffusion" title="Facilitated diffusion">facilitated diffusion</a> of protons into the matrix. The process results in the unharnessed potential energy of the proton <a href="/wiki/Electrochemistry" title="Electrochemistry">electrochemical</a> gradient being released as heat.<a href="#cite_note-Voet-2006-21">[21]</a> The process is mediated by a proton channel called <a href="/wiki/Thermogenin" title="Thermogenin">thermogenin</a>, or <a href="/wiki/UCP1" class="mw-redirect" title="UCP1">UCP1</a>.<a href="#cite_note-Mozo-2005-49">[49]</a> Thermogenin is primarily found in <a href="/wiki/Brown_adipose_tissue" title="Brown adipose tissue">brown adipose tissue</a>, or brown fat, and is responsible for non-shivering thermogenesis. Brown adipose tissue is found in mammals, and is at its highest levels in early life and in hibernating animals. In humans, brown adipose tissue is present at birth and decreases with age.<a href="#cite_note-Mozo-2005-49">[49]</a> <div class="mw-heading mw-heading3"><h3 id="Mitochondrial_fatty_acid_synthesis">Mitochondrial fatty acid synthesis</h3></div> Mitochondrial fatty acid synthesis (mtFASII) is essential for cellular respiration and mitochondrial biogenesis.<a href="#cite_note-50">[50]</a> It is also thought to play a role as a mediator in <a href="/wiki/Cell_signaling" title="Cell signaling">intracellular signaling</a> due to its influence on the levels of bioactive lipids, such as <a href="/wiki/Lysophospholipase" title="Lysophospholipase">lysophospholipids</a> and <a href="/wiki/Sphingolipid" title="Sphingolipid">sphingolipids</a>.<a href="#cite_note-51">[51]</a> <a href="/wiki/Octanoyl-(acyl-carrier_protein):protein_N-octanoyltransferase" class="mw-redirect" title="Octanoyl-(acyl-carrier protein):protein N-octanoyltransferase">Octanoyl-ACP</a> (C8) is considered to be the most important end product of mtFASII, which also forms the starting substrate of <a href="/wiki/Lipoic_acid" title="Lipoic acid">lipoic acid</a> biosynthesis.<a href="#cite_note-Nowinski-2018-52">[52]</a> Since lipoic acid is the cofactor of important mitochondrial enzyme complexes, such as the <a href="/wiki/Pyruvate_dehydrogenase_complex" title="Pyruvate dehydrogenase complex">pyruvate dehydrogenase complex</a> (PDC), <a href="/wiki/Oxoglutarate_dehydrogenase_complex" title="Oxoglutarate dehydrogenase complex">α-ketoglutarate dehydrogenase complex</a> (OGDC), <a href="/wiki/Branched-chain_alpha-keto_acid_dehydrogenase_complex" title="Branched-chain alpha-keto acid dehydrogenase complex">branched-chain α-ketoacid dehydrogenase complex</a> (BCKDC), and in the <a href="/wiki/Glycine_cleavage_system" title="Glycine cleavage system">glycine cleavage system</a> (GCS), mtFASII has an influence on energy metabolism.<a href="#cite_note-53">[53]</a> Other products of mtFASII play a role in the regulation of mitochondrial translation, <a href="/wiki/Iron-sulfur_cluster_biosynthesis" class="mw-redirect" title="Iron-sulfur cluster biosynthesis">FeS cluster biogenesis</a> and assembly of oxidative phosphorylation complexes.<a href="#cite_note-Nowinski-2018-52">[52]</a> Furthermore, with the help of mtFASII and acylated ACP, acetyl-CoA regulates its consumption in mitochondria.<a href="#cite_note-Nowinski-2018-52">[52]</a> <div class="mw-heading mw-heading3"><h3 id="Uptake,_storage_and_release_of_calcium_ions">Uptake, storage and release of calcium ions</h3></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Chondrocyte-_calcium_stain.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Chondrocyte-_calcium_stain.jpg/400px-Chondrocyte-_calcium_stain.jpg" decoding="async" width="400" height="204" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Chondrocyte-_calcium_stain.jpg/600px-Chondrocyte-_calcium_stain.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/8/83/Chondrocyte-_calcium_stain.jpg 2x" data-file-width="648" data-file-height="331" /></a><figcaption><a href="/wiki/Transmission_electron_microscope" class="mw-redirect" title="Transmission electron microscope">Transmission</a> <a href="/wiki/Micrograph" title="Micrograph">electron micrograph</a> of a <a href="/wiki/Chondrocyte" title="Chondrocyte">chondrocyte</a>, stained for calcium, showing its nucleus (N) and mitochondria (M)</figcaption></figure> The concentrations of free calcium in the cell can regulate an array of reactions and is important for <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> in the cell. Mitochondria can transiently <a href="/wiki/Calcium_storage" class="mw-redirect" title="Calcium storage">store calcium</a>, a contributing process for the cell's homeostasis of calcium.<a href="#cite_note-Santulli-2015c-54">[54]</a> <a href="#cite_note-Siegel-1999-55">[55]</a> Their ability to rapidly take in calcium for later release makes them good "cytosolic buffers" for calcium.<a href="#cite_note-Rossier-2006-56">[56]</a><a href="#cite_note-57">[57]</a><a href="#cite_note-58">[58]</a> The endoplasmic reticulum (ER) is the most significant storage site of calcium,<a href="#cite_note-Santulli-2015b-59">[59]</a> and there is a significant interplay between the mitochondrion and ER with regard to calcium.<a href="#cite_note-60">[60]</a> The calcium is taken up into the <a href="/wiki/Mitochondrial_matrix" title="Mitochondrial matrix">matrix</a> by the <a href="/wiki/Mitochondrial_calcium_uniporter" title="Mitochondrial calcium uniporter">mitochondrial calcium uniporter</a> on the <a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">inner mitochondrial membrane</a>.<a href="#cite_note-Miller-1998-61">[61]</a> It is primarily driven by the mitochondrial <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a>.<a href="#cite_note-Siegel-1999-55">[55]</a> Release of this calcium back into the cell's interior can occur via a sodium-calcium exchange protein or via "calcium-induced-calcium-release" pathways.<a href="#cite_note-Miller-1998-61">[61]</a> This can initiate calcium spikes or calcium waves with large changes in the membrane potential. These can activate a series of <a href="/wiki/Second_messenger_system" title="Second messenger system">second messenger system</a> proteins that can coordinate processes such as <a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">neurotransmitter release</a> in nerve cells and release of <a href="/wiki/Hormone" title="Hormone">hormones</a> in endocrine cells.<a href="#cite_note-Santulli-2015a-62">[62]</a> Ca2+ influx to the mitochondrial matrix has recently been implicated as a mechanism to regulate respiratory <a href="/wiki/Bioenergetics" title="Bioenergetics">bioenergetics</a> by allowing the electrochemical potential across the membrane to transiently "pulse" from ΔΨ-dominated to pH-dominated, facilitating a reduction of <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a>.<a href="#cite_note-63">[63]</a> In neurons, concomitant increases in cytosolic and mitochondrial calcium act to synchronize neuronal activity with mitochondrial energy metabolism. Mitochondrial matrix calcium levels can reach the tens of micromolar levels, which is necessary for the activation of <a href="/wiki/Isocitrate_dehydrogenase" title="Isocitrate dehydrogenase">isocitrate dehydrogenase</a>, one of the key regulatory enzymes of the <a href="/wiki/Krebs_cycle" class="mw-redirect" title="Krebs cycle">Krebs cycle</a>.<a href="#cite_note-64">[64]</a> <div class="mw-heading mw-heading3"><h3 id="Cellular_proliferation_regulation">Cellular proliferation regulation</h3></div> The relationship between cellular proliferation and mitochondria has been investigated. Tumor cells require ample ATP to synthesize bioactive compounds such as <a href="/wiki/Lipid" title="Lipid">lipids</a>, <a href="/wiki/Protein" title="Protein">proteins</a>, and <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a> for rapid proliferation.<a href="#cite_note-Weinberg-2009-65">[65]</a> The majority of ATP in tumor cells is generated via the <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a> pathway (OxPhos).<a href="#cite_note-Moreno-Sánchez-2007-66">[66]</a> Interference with OxPhos cause <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> arrest suggesting that mitochondria play a role in cell proliferation.<a href="#cite_note-Moreno-Sánchez-2007-66">[66]</a> Mitochondrial ATP production is also vital for <a href="/wiki/Cell_division" title="Cell division">cell division</a> and differentiation in infection<a href="#cite_note-67">[67]</a> in addition to basic functions in the cell including the regulation of cell volume, solute <a href="/wiki/Concentration" title="Concentration">concentration</a>, and cellular architecture.<a href="#cite_note-Pedersen-1994-68">[68]</a><a href="#cite_note-Pattappa-2011-69">[69]</a><a href="#cite_note-Agarwal-2011-70">[70]</a> ATP levels differ at various stages of the cell cycle suggesting that there is a relationship between the abundance of ATP and the cell's ability to enter a new cell cycle.<a href="#cite_note-Sweet-1999-71">[71]</a> ATP's role in the basic functions of the cell make the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> sensitive to changes in the availability of mitochondrial derived ATP.<a href="#cite_note-Sweet-1999-71">[71]</a> The variation in ATP levels at different stages of the cell cycle support the hypothesis that mitochondria play an important role in cell cycle regulation.<a href="#cite_note-Sweet-1999-71">[71]</a> Although the specific mechanisms between mitochondria and the cell cycle regulation is not well understood, studies have shown that low energy cell cycle checkpoints monitor the energy capability before committing to another round of cell division.<a href="#cite_note-McBride-2006-12">[12]</a> <div class="mw-heading mw-heading3"><h3 id="Programmed_cell_death_and_innate_immunity">Programmed cell death and innate immunity</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Programmed_cell_death#Intrinsic_Pathway" title="Programmed cell death">Programmed cell death § Intrinsic Pathway</a></div> <a href="/wiki/Programmed_cell_death" title="Programmed cell death">Programmed cell death</a> (PCD) is crucial for various physiological functions, including organ development and cellular homeostasis. It serves as an intrinsic mechanism to prevent <a href="/wiki/Malignant_transformation" title="Malignant transformation">malignant transformation</a> and plays a fundamental role in <a href="/wiki/Immune_system" title="Immune system">immunity</a> by aiding in antiviral defense, pathogen elimination, inflammation, and immune cell recruitment.<a href="#cite_note-72">[72]</a> Mitochondria have long been recognized for their central role in the <a href="/wiki/Apoptosis#Intrinsic_pathway" title="Apoptosis">intrinsic pathway</a> of <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, a form of PCD.<a href="#cite_note-73">[73]</a> In recent decades, they have also been identified as a signalling hub for much of the <a href="/wiki/Innate_immune_system" title="Innate immune system">innate immune system</a>.<a href="#cite_note-Bahat-2021-74">[74]</a> The <a href="/wiki/Symbiogenesis" title="Symbiogenesis">endosymbiotic origin</a> of mitochondria distinguishes them from other cellular components, and the exposure of mitochondrial elements to the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> can trigger the same pathways as infection markers. These pathways lead to <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, <a href="/wiki/Autophagy" title="Autophagy">autophagy</a>, or the induction of proinflammatory genes.<a href="#cite_note-Murphy-2024-75">[75]</a><a href="#cite_note-Bahat-2021-74">[74]</a> Mitochondria contribute to apoptosis by releasing <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome c</a>, which directly induces the formation of <a href="/wiki/Apoptosome" title="Apoptosome">apoptosomes</a>. Additionally, they are a source of various <a href="/wiki/Damage-associated_molecular_pattern" title="Damage-associated molecular pattern">damage-associated molecular patterns</a> (DAMPs). These DAMPs are often recognised by the same <a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern-recognition receptors</a> (PRRs) that respond to <a href="/wiki/Pathogen-associated_molecular_pattern" title="Pathogen-associated molecular pattern">pathogen-associated molecular patterns</a> (PAMPs) during infections.<a href="#cite_note-76">[76]</a> For example, mitochondrial mtDNA resembles bacterial DNA due to its lack of <a href="/wiki/CpG_site" title="CpG site">CpG</a> methylation and can be detected by <a href="/wiki/Toll-like_receptor_9" title="Toll-like receptor 9">Toll-like receptor 9</a> and <a href="/wiki/CGAS%E2%80%93STING_cytosolic_DNA_sensing_pathway" title="CGAS–STING cytosolic DNA sensing pathway">cGAS</a>.<a href="#cite_note-77">[77]</a> <a href="/wiki/DsRNA" class="mw-redirect" title="DsRNA">Double-stranded RNA</a> (dsRNA), produced due to bidirectional mitochondrial transcription, can activate viral sensing pathways through <a href="/wiki/RIG-I-like_receptor" title="RIG-I-like receptor">RIG-I-like receptors</a>.<a href="#cite_note-78">[78]</a> Additionally, the N-formylation of <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">mitochondrial proteins</a>, similar to that of bacterial proteins, can be recognized by <a href="/wiki/Formyl_peptide_receptor" title="Formyl peptide receptor">formyl peptide receptors</a>.<a href="#cite_note-79">[79]</a><a href="#cite_note-80">[80]</a> Normally, these mitochondrial components are sequestered from the rest of the cell but are released following mitochondrial membrane permeabilization during apoptosis or passively after mitochondrial damage. However, mitochondria also play an active role in innate immunity, releasing mtDNA in response to metabolic cues.<a href="#cite_note-Bahat-2021-74">[74]</a> Mitochondria are also the <a href="/wiki/Subcellular_localization" title="Subcellular localization">localization site</a> for immune and apoptosis regulatory proteins, such as <a href="/wiki/Apoptosis_regulator_BAX" title="Apoptosis regulator BAX">BAX</a>, <a href="/wiki/Mitochondrial_antiviral-signaling_protein" title="Mitochondrial antiviral-signaling protein">MAVS</a> (located on the <a class="mw-selflink-fragment" href="#Outer_membrane">outer membrane</a>), and <a href="/wiki/NLRX1" title="NLRX1">NLRX1</a> (found in the <a class="mw-selflink-fragment" href="#Matrix">matrix</a>). These proteins are modulated by the mitochondrial metabolic status and mitochondrial dynamics.<a href="#cite_note-Bahat-2021-74">[74]</a><a href="#cite_note-81">[81]</a><a href="#cite_note-82">[82]</a> <div class="mw-heading mw-heading3"><h3 id="Additional_functions">Additional functions</h3></div> Mitochondria play a central role in many other <a href="/wiki/Metabolism" title="Metabolism">metabolic</a> tasks, such as: <ul><li>Signaling through mitochondrial <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a><a href="#cite_note-Li-2013-83">[83]</a></li> <li>Regulation of the <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a><a href="#cite_note-Voet-2006-21">[21]</a></li> <li>Calcium signaling (including calcium-evoked apoptosis)<a href="#cite_note-84">[84]</a></li> <li>Regulation of cellular <a href="/wiki/Metabolism" title="Metabolism">metabolism</a><a href="#cite_note-McBride-2006-12">[12]</a></li> <li>Certain <a href="/wiki/Heme" title="Heme">heme</a> synthesis reactions<a href="#cite_note-85">[85]</a> (see also: <a href="/wiki/Porphyrin" title="Porphyrin">Porphyrin</a>)</li> <li><a href="/wiki/Steroid" title="Steroid">Steroid</a> synthesis<a href="#cite_note-Rossier-2006-56">[56]</a></li> <li>Hormonal signaling<a href="#cite_note-86">[86]</a> – mitochondria are sensitive and responsive to hormones, in part by the action of mitochondrial estrogen receptors (mtERs). These receptors have been found in various tissues and cell types, including brain<a href="#cite_note-87">[87]</a> and heart<a href="#cite_note-88">[88]</a></li> <li>Development and function of immune cells<a href="#cite_note-89">[89]</a></li> <li>Neuronal mitochondria also contribute to cellular quality control by reporting neuronal status towards microglia through specialised somatic-junctions.<a href="#cite_note-Cserép-2020-90">[90]</a></li> <li>Mitochondria of developing neurons contribute to intercellular signaling towards <a href="/wiki/Microglia" title="Microglia">microglia</a>, which communication is indispensable for proper regulation of brain development.<a href="#cite_note-91">[91]</a></li></ul> Some mitochondrial functions are performed only in specific types of cells. For example, mitochondria in <a href="/wiki/Liver_cell" class="mw-redirect" title="Liver cell">liver cells</a> contain enzymes that allow them to detoxify <a href="/wiki/Ammonia" title="Ammonia">ammonia</a>, a waste product of protein metabolism. A mutation in the genes regulating any of these functions can result in <a href="/wiki/Mitochondrial_disease" title="Mitochondrial disease">mitochondrial diseases</a>. Mitochondrial proteins (proteins transcribed from mitochondrial DNA) vary depending on the tissue and the species. In humans, 615 distinct types of proteins have been identified from <a href="/wiki/Heart" title="Heart">cardiac</a> mitochondria,<a href="#cite_note-92">[92]</a> whereas in <a href="/wiki/Murinae" title="Murinae">rats</a>, 940 proteins have been reported.<a href="#cite_note-93">[93]</a> The mitochondrial <a href="/wiki/Proteome" title="Proteome">proteome</a> is thought to be dynamically regulated.<a href="#cite_note-94">[94]</a> <div class="mw-heading mw-heading2"><h2 id="Organization_and_distribution">Organization and distribution</h2></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:HeLa_mtGFP.tif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/HeLa_mtGFP.tif/lossy-page1-220px-HeLa_mtGFP.tif.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/90/HeLa_mtGFP.tif/lossy-page1-330px-HeLa_mtGFP.tif.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/90/HeLa_mtGFP.tif/lossy-page1-440px-HeLa_mtGFP.tif.jpg 2x" data-file-width="1024" data-file-height="1024" /></a><figcaption>Typical mitochondrial network (green) in two human cells (<a href="/wiki/HeLa_cells" class="mw-redirect" title="HeLa cells">HeLa cells</a>)</figcaption></figure> Mitochondria (or related structures) are found in all <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> (except the <a href="/wiki/Oxymonad" title="Oxymonad">Oxymonad</a> <a href="/wiki/Monocercomonoides" title="Monocercomonoides">Monocercomonoides</a>).<a href="#cite_note-Karnkowska-2016-5">[5]</a> Although commonly depicted as bean-like structures they form a highly dynamic network in the majority of cells where they constantly undergo <a href="/wiki/Mitochondrial_fission" title="Mitochondrial fission">fission</a> and <a href="/wiki/Mitochondrial_fusion" title="Mitochondrial fusion">fusion</a>. The population of all the mitochondria of a given cell constitutes the chondriome.<a href="#cite_note-Logan-2010-95">[95]</a> Mitochondria vary in number and location according to cell type. A single mitochondrion is often found in unicellular organisms, while human liver cells have about 1000–2000 mitochondria per cell, making up 1/5 of the cell volume.<a href="#cite_note-Alberts-2005-20">[20]</a> The mitochondrial content of otherwise similar cells can vary substantially in size and membrane potential,<a href="#cite_note-96">[96]</a> with differences arising from sources including uneven partitioning at cell division, leading to <a href="/wiki/Cellular_noise" title="Cellular noise">extrinsic differences</a> in ATP levels and downstream cellular processes.<a href="#cite_note-97">[97]</a> The mitochondria can be found nestled between <a href="/wiki/Myofibril" title="Myofibril">myofibrils</a> of <a href="/wiki/Muscle" title="Muscle">muscle</a> or wrapped around the <a href="/wiki/Sperm" title="Sperm">sperm</a> <a href="/wiki/Flagellum" title="Flagellum">flagellum</a>.<a href="#cite_note-Alberts-2005-20">[20]</a> Often, they form a complex 3D branching network inside the cell with the <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a>. The association with the cytoskeleton determines mitochondrial shape, which can affect the function as well:<a href="#cite_note-98">[98]</a> different structures of the mitochondrial network may afford the population a variety of physical, chemical, and signalling advantages or disadvantages.<a href="#cite_note-99">[99]</a> Mitochondria in cells are always distributed along microtubules and the distribution of these organelles is also correlated with the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>.<a href="#cite_note-100">[100]</a> Recent evidence suggests that <a href="/wiki/Vimentin" title="Vimentin">vimentin</a>, one of the components of the cytoskeleton, is also critical to the association with the cytoskeleton.<a href="#cite_note-101">[101]</a> <div class="mw-heading mw-heading3"><h3 id="Mitochondria-associated_ER_membrane_(MAM)">Mitochondria-associated ER membrane (MAM)</h3></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/Mitochondria_associated_membranes" title="Mitochondria associated membranes">Mitochondria associated membranes</a></div> The mitochondria-associated ER membrane (MAM) is another structural element that is increasingly recognized for its critical role in cellular physiology and <a href="/wiki/Homeostasis" title="Homeostasis">homeostasis</a>. Once considered a technical snag in cell fractionation techniques, the alleged ER vesicle contaminants that invariably appeared in the mitochondrial fraction have been re-identified as membranous structures derived from the MAM—the interface between mitochondria and the ER.<a href="#cite_note-Rizzuto-2009-102">[102]</a> Physical coupling between these two organelles had previously been observed in electron micrographs and has more recently been probed with <a href="/wiki/Fluorescence_microscopy" class="mw-redirect" title="Fluorescence microscopy">fluorescence microscopy</a>.<a href="#cite_note-Rizzuto-2009-102">[102]</a> Such studies estimate that at the MAM, which may comprise up to 20% of the mitochondrial outer membrane, the ER and mitochondria are separated by a mere 10–25 nm and held together by protein tethering complexes.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Hayashi-2009-30">[30]</a><a href="#cite_note-de_Brito-2010-103">[103]</a> Purified MAM from subcellular fractionation is enriched in enzymes involved in phospholipid exchange, in addition to channels associated with Ca2+ signaling.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-de_Brito-2010-103">[103]</a> These hints of a prominent role for the MAM in the regulation of cellular lipid stores and signal transduction have been borne out, with significant implications for mitochondrial-associated cellular phenomena, as discussed below. Not only has the MAM provided insight into the mechanistic basis underlying such physiological processes as intrinsic apoptosis and the propagation of calcium signaling, but it also favors a more refined view of the mitochondria. Though often seen as static, isolated 'powerhouses' hijacked for cellular metabolism through an ancient endosymbiotic event, the evolution of the MAM underscores the extent to which mitochondria have been integrated into overall cellular physiology, with intimate physical and functional coupling to the endomembrane system. <div class="mw-heading mw-heading4"><h4 id="Phospholipid_transfer">Phospholipid transfer</h4></div> The MAM is enriched in enzymes involved in lipid biosynthesis, such as phosphatidylserine synthase on the ER face and phosphatidylserine decarboxylase on the mitochondrial face.<a href="#cite_note-Vance-1996-104">[104]</a><a href="#cite_note-Lebiedzinska-2009-105">[105]</a> Because mitochondria are dynamic organelles constantly undergoing <a href="/wiki/Mitochondrial_fission" title="Mitochondrial fission">fission</a> and <a href="/wiki/Mitochondrial_fusion" title="Mitochondrial fusion">fusion</a> events, they require a constant and well-regulated supply of phospholipids for membrane integrity.<a href="#cite_note-Twig-2008-106">[106]</a><a href="#cite_note-Osman-2011-107">[107]</a> But mitochondria are not only a destination for the phospholipids they finish synthesis of; rather, this organelle also plays a role in inter-organelle trafficking of the intermediates and products of phospholipid biosynthetic pathways, ceramide and cholesterol metabolism, and glycosphingolipid anabolism.<a href="#cite_note-Lebiedzinska-2009-105">[105]</a><a href="#cite_note-Osman-2011-107">[107]</a> Such trafficking capacity depends on the MAM, which has been shown to facilitate transfer of lipid intermediates between organelles.<a href="#cite_note-Vance-1996-104">[104]</a> In contrast to the standard vesicular mechanism of lipid transfer, evidence indicates that the physical proximity of the ER and mitochondrial membranes at the MAM allows for lipid flipping between opposed bilayers.<a href="#cite_note-Osman-2011-107">[107]</a> Despite this unusual and seemingly energetically unfavorable mechanism, such transport does not require ATP.<a href="#cite_note-Osman-2011-107">[107]</a> Instead, in yeast, it has been shown to be dependent on a <a href="/wiki/Protein_complex" title="Protein complex">multiprotein</a> tethering structure termed the ER-mitochondria encounter structure, or ERMES, although it remains unclear whether this structure directly mediates lipid transfer or is required to keep the membranes in sufficiently close proximity to lower the <a href="/wiki/Activation_energy" title="Activation energy">energy barrier</a> for <a href="/wiki/Lipid" title="Lipid">lipid</a> flipping.<a href="#cite_note-Osman-2011-107">[107]</a><a href="#cite_note-Kornmann-2009-108">[108]</a> The MAM may also be part of the secretory pathway, in addition to its role in intracellular lipid trafficking. In particular, the MAM appears to be an intermediate destination between the rough ER and the Golgi in the pathway that leads to <a href="/wiki/Very-low-density_lipoprotein" class="mw-redirect" title="Very-low-density lipoprotein">very-low-density lipoprotein</a>, or VLDL, assembly and secretion.<a href="#cite_note-Lebiedzinska-2009-105">[105]</a><a href="#cite_note-Rusiñol-1994-109">[109]</a> The MAM thus serves as a critical metabolic and trafficking hub in lipid metabolism. <div class="mw-heading mw-heading4"><h4 id="Calcium_signaling">Calcium signaling</h4></div> A critical role for the ER in calcium signaling was acknowledged before such a role for the mitochondria was widely accepted, in part because the low affinity of Ca2+ channels localized to the outer mitochondrial membrane seemed to contradict this organelle's purported responsiveness to changes in intracellular Ca2+ flux.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Santulli-2015b-59">[59]</a> But the presence of the MAM resolves this apparent contradiction: the close physical association between the two organelles results in Ca2+ microdomains at contact points that facilitate efficient Ca2+ transmission from the ER to the mitochondria.<a href="#cite_note-Rizzuto-2009-102">[102]</a> Transmission occurs in response to so-called "Ca2+ puffs" generated by spontaneous clustering and activation of <a href="/wiki/IP3R" class="mw-redirect" title="IP3R">IP3R</a>, a canonical ER membrane Ca2+ channel.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Hayashi-2009-30">[30]</a> The fate of these puffs—in particular, whether they remain restricted to isolated locales or integrated into Ca2+ waves for propagation throughout the cell—is determined in large part by MAM dynamics. Although reuptake of Ca2+ by the ER (concomitant with its release) modulates the intensity of the puffs, thus insulating mitochondria to a certain degree from high Ca2+ exposure, the MAM often serves as a firewall that essentially buffers Ca2+ puffs by acting as a sink into which free ions released into the cytosol can be funneled.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Kopach-2008-110">[110]</a><a href="#cite_note-Csordás-2001-111">[111]</a> This Ca2+ tunneling occurs through the low-affinity Ca2+ receptor <a href="/wiki/VDAC1" title="VDAC1">VDAC1</a>, which recently has been shown to be physically <a href="/wiki/Tether_(cell_biology)" title="Tether (cell biology)">tethered</a> to the IP3R clusters on the ER membrane and enriched at the MAM.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Hayashi-2009-30">[30]</a><a href="#cite_note-Decuypere-2011-112">[112]</a> The ability of mitochondria to serve as a Ca2+ sink is a result of the electrochemical gradient generated during oxidative phosphorylation, which makes tunneling of the cation an exergonic process.<a href="#cite_note-Decuypere-2011-112">[112]</a> Normal, mild calcium influx from cytosol into the mitochondrial matrix causes transient depolarization that is corrected by pumping out protons. But transmission of Ca2+ is not unidirectional; rather, it is a two-way street.<a href="#cite_note-Santulli-2015b-59">[59]</a> The properties of the Ca2+ pump SERCA and the channel IP3R present on the ER membrane facilitate feedback regulation coordinated by MAM function. In particular, the clearance of Ca2+ by the MAM allows for <a href="/wiki/Spatio-temporal_pattern" class="mw-redirect" title="Spatio-temporal pattern">spatio-temporal patterning</a> of Ca2+ signaling because Ca2+ alters IP3R activity in a biphasic manner.<a href="#cite_note-Rizzuto-2009-102">[102]</a> <a href="/wiki/SERCA" title="SERCA">SERCA</a> is likewise affected by mitochondrial feedback: uptake of Ca2+ by the MAM stimulates ATP production, thus providing energy that enables SERCA to reload the ER with Ca2+ for continued Ca2+ efflux at the MAM.<a href="#cite_note-Kopach-2008-110">[110]</a><a href="#cite_note-Decuypere-2011-112">[112]</a> Thus, the MAM is not a passive buffer for Ca2+ puffs; rather it helps modulate further Ca2+ signaling through feedback loops that affect ER dynamics. Regulating ER release of Ca2+ at the MAM is especially critical because only a certain window of Ca2+ uptake sustains the mitochondria, and consequently the cell, at homeostasis. Sufficient intraorganelle Ca2+ signaling is required to stimulate metabolism by activating dehydrogenase enzymes critical to flux through the citric acid cycle.<a href="#cite_note-Diercks-2017-113">[113]</a><a href="#cite_note-Hajnóczky-2011-114">[114]</a> However, once Ca2+ signaling in the mitochondria passes a certain threshold, it stimulates the intrinsic pathway of apoptosis in part by collapsing the mitochondrial membrane potential required for metabolism.<a href="#cite_note-Rizzuto-2009-102">[102]</a> Studies examining the role of pro- and anti-apoptotic factors support this model; for example, the anti-apoptotic factor Bcl-2 has been shown to interact with IP3Rs to reduce Ca2+ filling of the ER, leading to reduced efflux at the MAM and preventing collapse of the mitochondrial membrane potential post-apoptotic stimuli.<a href="#cite_note-Rizzuto-2009-102">[102]</a> Given the need for such fine regulation of Ca2+ signaling, it is perhaps unsurprising that dysregulated mitochondrial Ca2+ has been implicated in several neurodegenerative diseases, while the catalogue of tumor suppressors includes a few that are enriched at the MAM.<a href="#cite_note-Decuypere-2011-112">[112]</a> <div class="mw-heading mw-heading4"><h4 id="Molecular_basis_for_tethering">Molecular basis for tethering</h4></div> Recent advances in the identification of the <a href="/wiki/Tether_(cell_biology)" title="Tether (cell biology)">tethers</a> between the mitochondrial and ER membranes suggest that the scaffolding function of the molecular elements involved is secondary to other, non-structural functions. In yeast, ERMES, a multiprotein complex of interacting ER- and mitochondrial-resident membrane proteins, is required for lipid transfer at the MAM and exemplifies this principle. One of its components, for example, is also a constituent of the protein complex required for insertion of transmembrane beta-barrel proteins into the lipid bilayer.<a href="#cite_note-Osman-2011-107">[107]</a> However, a <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologue</a> of the ERMES complex has not yet been identified in mammalian cells. Other proteins implicated in scaffolding likewise have functions independent of structural tethering at the MAM; for example, ER-resident and mitochondrial-resident mitofusins form heterocomplexes that regulate the number of inter-organelle contact sites, although mitofusins were first identified for their role in <a href="/wiki/Mitochondrial_fission" title="Mitochondrial fission">fission</a> and <a href="/wiki/Mitochondrial_fusion" title="Mitochondrial fusion">fusion</a> events between individual mitochondria.<a href="#cite_note-Rizzuto-2009-102">[102]</a> <a href="/wiki/Glucose" title="Glucose">Glucose</a>-related protein 75 (grp75) is another dual-function protein. In addition to the matrix pool of grp75, a portion serves as a chaperone that physically links the mitochondrial and ER Ca2+ channels VDAC and IP3R for efficient Ca2+ transmission at the MAM.<a href="#cite_note-Rizzuto-2009-102">[102]</a><a href="#cite_note-Hayashi-2009-30">[30]</a> Another potential tether is <a href="/wiki/Sigma-1_receptor" title="Sigma-1 receptor">Sigma-1R</a>, a non-opioid receptor whose stabilization of ER-resident IP3R may preserve communication at the MAM during the metabolic stress response.<a href="#cite_note-115">[115]</a><a href="#cite_note-116">[116]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:ERMES.png" class="mw-file-description"><img alt="ERMES tethering complex." src="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/ERMES.png/220px-ERMES.png" decoding="async" width="220" height="226" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/b/ba/ERMES.png 1.5x" data-file-width="230" data-file-height="236" /></a><figcaption>Model of the yeast multimeric tethering complex, ERMES</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Perspective">Perspective</h4></div> The MAM is a critical signaling, metabolic, and trafficking hub in the cell that allows for the integration of ER and mitochondrial physiology. Coupling between these organelles is not simply structural but functional as well and critical for overall cellular physiology and <a href="/wiki/Homeostasis" title="Homeostasis">homeostasis</a>. The MAM thus offers a perspective on mitochondria that diverges from the traditional view of this organelle as a static, isolated unit appropriated for its metabolic capacity by the cell.<a href="#cite_note-Csordás-2018-117">[117]</a> Instead, this mitochondrial-ER interface emphasizes the integration of the mitochondria, the product of an endosymbiotic event, into diverse cellular processes. Recently it has also been shown, that mitochondria and MAM-s in neurons are anchored to specialised intercellular communication sites (so called somatic-junctions). <a href="/wiki/Microglia" title="Microglia">Microglial</a> processes monitor and protect neuronal functions at these sites, and MAM-s are supposed to have an important role in this type of cellular quality-control.<a href="#cite_note-Cserép-2020-90">[90]</a> <div class="mw-heading mw-heading2"><h2 id="Origin_and_evolution">Origin and evolution</h2></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/Symbiogenesis" title="Symbiogenesis">Symbiogenesis</a></div> There are two hypotheses about the origin of mitochondria: <a href="/wiki/Endosymbiotic_theory" class="mw-redirect" title="Endosymbiotic theory">endosymbiotic</a> and <a href="/wiki/Autotransplantation" title="Autotransplantation">autogenous</a>. The endosymbiotic hypothesis suggests that mitochondria were originally <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotic</a> cells, capable of implementing oxidative mechanisms that were not possible for eukaryotic cells; they became <a href="/wiki/Endosymbiont" title="Endosymbiont">endosymbionts</a> living inside the eukaryote.<a href="#cite_note-Gabaldón-2021-23">[23]</a><a href="#cite_note-McCutcheon-2021-118">[118]</a><a href="#cite_note-Callier-2022-119">[119]</a><a href="#cite_note-Margulis-1986-120">[120]</a> In the autogenous hypothesis, mitochondria were born by splitting off a portion of DNA from the nucleus of the eukaryotic cell at the time of divergence with the prokaryotes; this DNA portion would have been enclosed by membranes, which could not be crossed by proteins. Since mitochondria have many features in common with <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, the endosymbiotic hypothesis is the more widely accepted of the two accounts.<a href="#cite_note-Margulis-1986-120">[120]</a><a href="#cite_note-121">[121]</a> A <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrion contains DNA</a>, which is organized as several copies of a single, usually <a href="/wiki/Mitochondrial_DNA#Circular_versus_linear" title="Mitochondrial DNA">circular</a> <a href="/wiki/Chromosome" title="Chromosome">chromosome</a>. This mitochondrial chromosome contains genes for <a href="/wiki/Redox" title="Redox">redox</a> proteins, such as those of the respiratory chain. The <a href="/wiki/CoRR_hypothesis" title="CoRR hypothesis">CoRR hypothesis</a> proposes that this co-location is required for redox regulation. The mitochondrial <a href="/wiki/Genome" title="Genome">genome</a> codes for some <a href="/wiki/RNA" title="RNA">RNAs</a> of <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>, and the 22 <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNAs</a> necessary for the translation of <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNAs</a> into protein. The circular structure is also found in prokaryotes. The <a href="/wiki/Proto-mitochondrion" title="Proto-mitochondrion">proto-mitochondrion</a> was probably closely related to <a href="/wiki/Rickettsia" title="Rickettsia">Rickettsia</a>.<a href="#cite_note-122">[122]</a><a href="#cite_note-123">[123]</a> However, the exact relationship of the ancestor of mitochondria to the <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">alphaproteobacteria</a> and whether the mitochondrion was formed at the same time or after the nucleus, remains controversial.<a href="#cite_note-124">[124]</a> For example, it has been suggested that the <a href="/wiki/SAR11_clade" class="mw-redirect" title="SAR11 clade">SAR11 clade</a> of bacteria shares a relatively recent common ancestor with the mitochondria,<a href="#cite_note-125">[125]</a> while <a href="/wiki/Phylogenomic" class="mw-redirect" title="Phylogenomic">phylogenomic</a> analyses indicate that mitochondria evolved from a <a href="/wiki/Pseudomonadota" title="Pseudomonadota">Pseudomonadota</a> lineage that is closely related to or a member of <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">alphaproteobacteria</a>.<a href="#cite_note-Martijn-2018-126">[126]</a><a href="#cite_note-127">[127]</a> Some papers describe mitochondria as sister to the alphaproteobactera, together forming the sister the marineproteo1 group, together forming the sister to <a href="/wiki/Magnetococcidae" class="mw-redirect" title="Magnetococcidae">Magnetococcidae</a>.<a href="#cite_note-128">[128]</a><a href="#cite_note-129">[129]</a><a href="#cite_note-130">[130]</a><a href="#cite_note-131">[131]</a> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1258728058">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%;line-height:inherit}.mw-parser-output table.clade td.clade-label{min-width:0.2em;width:0.2em;padding:0.1em 0.25em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-label::before,.mw-parser-output table.clade td.clade-slabel::before{content:"\2060 "}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0.1em 0.25em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade" style="font-size:75%;line-height:75%"> <tbody><tr> <td class="clade-label first"><a href="/wiki/Proteobacteria" class="mw-redirect" title="Proteobacteria">Proteobacteria</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first" style="border-left:none;border-right:none;border-bottom:3px double;"> </td> <td rowspan="2" class="clade-leaf"> </td></tr> <tr> <td class="clade-slabel" style="border-right:none;border-left:1px solid;"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">Alphaproteobacteria s.l.</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Magnetococcales" title="Magnetococcales">Magnetococcales</a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/w/index.php?title=MarineProteo1&action=edit&redlink=1" class="new" title="MarineProteo1 (page does not exist)">MarineProteo1</a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Mitochondria </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">Alphaproteobacteria s.s.</a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> The ribosomes coded for by the mitochondrial DNA are similar to those from bacteria in size and structure.<a href="#cite_note-O'Brien-2003-132">[132]</a> They closely resemble the bacterial <a href="/wiki/Ribosome#Structure" title="Ribosome">70S</a> ribosome and not the <a href="/wiki/Ribosome#Structure" title="Ribosome">80S</a> <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasmic</a> ribosomes, which are coded for by <a href="/wiki/Cell_nucleus" title="Cell nucleus">nuclear</a> DNA. The <a href="/wiki/Endosymbiotic" class="mw-redirect" title="Endosymbiotic">endosymbiotic</a> relationship of mitochondria with their host cells was popularized by <a href="/wiki/Lynn_Margulis" title="Lynn Margulis">Lynn Margulis</a>.<a href="#cite_note-133">[133]</a> The <a href="/wiki/Endosymbiotic_theory" class="mw-redirect" title="Endosymbiotic theory">endosymbiotic hypothesis</a> suggests that mitochondria descended from aerobic bacteria that somehow survived <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a> by another cell, and became incorporated into the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. The ability of these bacteria to conduct <a href="/wiki/Cellular_respiration" title="Cellular respiration">respiration</a> in host cells that had relied on <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a> and <a href="/wiki/Fermentation_(biochemistry)" class="mw-redirect" title="Fermentation (biochemistry)">fermentation</a> would have provided a considerable evolutionary advantage. This symbiotic relationship probably developed 1.7 to 2 billion years ago.<a href="#cite_note-134">[134]</a><a href="#cite_note-135">[135]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png/220px-The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png" decoding="async" width="220" height="151" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png/330px-The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/29/The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png/440px-The-origins-of-mitochondrion-related-organelles-A-hypothetical-scenario-for-the.png 2x" data-file-width="839" data-file-height="577" /></a><figcaption>Evolution of MROs</figcaption></figure> A few groups of unicellular eukaryotes have only vestigial mitochondria or derived structures: The <a href="/wiki/Microsporidia" title="Microsporidia">microsporidians</a>, <a href="/wiki/Metamonad" title="Metamonad">metamonads</a>, and <a href="/wiki/Archamoebae" title="Archamoebae">archamoebae</a>.<a href="#cite_note-Cavalier-Smith-1991-136">[136]</a> These groups appear as the most primitive eukaryotes on <a href="/wiki/Phylogenetic_trees" class="mw-redirect" title="Phylogenetic trees">phylogenetic trees</a> constructed using <a href="/wiki/RRNA" class="mw-redirect" title="RRNA">rRNA</a> information, which once suggested that they appeared before the origin of mitochondria. However, this is now known to be an artifact of <a href="/wiki/Long-branch_attraction" class="mw-redirect" title="Long-branch attraction">long-branch attraction</a>: They are derived groups and retain genes or organelles derived from mitochondria (e. g., <a href="/wiki/Mitosome" title="Mitosome">mitosomes</a> and <a href="/wiki/Hydrogenosome" title="Hydrogenosome">hydrogenosomes</a>).<a href="#cite_note-Henze-2003-8">[8]</a> Hydrogenosomes, mitosomes, and related organelles as found in some <a href="/wiki/Loricifera" title="Loricifera">loricifera</a> (e. g. <a href="/wiki/Spinoloricus" title="Spinoloricus">Spinoloricus</a>)<a href="#cite_note-137">[137]</a><a href="#cite_note-138">[138]</a> and <a href="/wiki/Myxozoa" title="Myxozoa">myxozoa</a> (e. g. <a href="/wiki/Henneguya_zschokkei" title="Henneguya zschokkei">Henneguya zschokkei</a>) are together classified as MROs, mitochondrion-related organelles.<a href="#cite_note-Yahalomi-2020-7">[7]</a><a href="#cite_note-Shiflett-2010-139">[139]</a> <a href="/wiki/Monocercomonoides" title="Monocercomonoides">Monocercomonoides</a> and other <a href="/wiki/Oxymonad" title="Oxymonad">oxymonads</a> appear to have lost their mitochondria completely and at least some of the mitochondrial functions seem to be carried out by cytoplasmic proteins now.<a href="#cite_note-Karnkowska-2016-5">[5]</a><a href="#cite_note-140">[140]</a><a href="#cite_note-Novák-2023-10">[10]</a> <div class="mw-heading mw-heading2"><h2 id="Mitochondrial_genetics">Mitochondrial genetics</h2></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Map_of_the_human_mitochondrial_genome.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Map_of_the_human_mitochondrial_genome.svg/300px-Map_of_the_human_mitochondrial_genome.svg.png" decoding="async" width="300" height="286" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Map_of_the_human_mitochondrial_genome.svg/450px-Map_of_the_human_mitochondrial_genome.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/15/Map_of_the_human_mitochondrial_genome.svg/600px-Map_of_the_human_mitochondrial_genome.svg.png 2x" data-file-width="1040" data-file-height="991" /></a><figcaption>The <a href="/wiki/Mitochondrial_DNA#Circular_versus_linear" title="Mitochondrial DNA">circular</a> 16,569 bp human mitochondrial genome encoding 37 genes, i.e., 28 on the H-strand and 9 on the L-strand</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial DNA</a></div> Mitochondria contain their own genome. The <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">human mitochondrial genome</a> is a circular double-stranded <a href="/wiki/DNA" title="DNA">DNA</a> molecule of about 16 <a href="/wiki/Base_pair" title="Base pair">kilobases</a>.<a href="#cite_note-Chan-2006-141">[141]</a> It encodes 37 genes: 13 for <a href="/wiki/Protein_subunit" title="Protein subunit">subunits</a> of respiratory complexes I, III, IV and V, 22 for mitochondrial <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a> (for the 20 standard amino acids, plus an extra gene for leucine and serine), and 2 for <a href="/wiki/RRNA" class="mw-redirect" title="RRNA">rRNA</a> (12S and 16S rRNA).<a href="#cite_note-Chan-2006-141">[141]</a> One mitochondrion can contain two to ten copies of its DNA.<a href="#cite_note-Wiesner-1992-142">[142]</a> One of the two mitochondrial DNA (mtDNA) strands has a disproportionately higher ratio of the heavier nucleotides adenine and guanine, and this is termed the heavy strand (or H strand), whereas the other strand is termed the light strand (or L strand). The weight difference allows the two strands to be separated by <a href="/wiki/Centrifugation" title="Centrifugation">centrifugation</a>. mtDNA has one long non-coding stretch known as the non-coding region (NCR), which contains the heavy strand promoter (HSP) and light strand promoter (LSP) for RNA transcription, the origin of replication for the H strand (OriH) localized on the L strand, three conserved sequence boxes (CSBs 1–3), and a termination-associated sequence (TAS). The origin of replication for the L strand (OriL) is localized on the H strand 11,000 bp downstream of OriH, located within a cluster of genes coding for tRNA.<a href="#cite_note-143">[143]</a> As in prokaryotes, there is a very high proportion of coding DNA and an absence of repeats. Mitochondrial genes are <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a> as multigenic transcripts, which are cleaved and <a href="/wiki/Polyadenylation" title="Polyadenylation">polyadenylated</a> to yield mature <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNAs</a>. Most proteins necessary for mitochondrial function are encoded by genes in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a> and the corresponding proteins are imported into the mitochondrion.<a href="#cite_note-Anderson-1981-144">[144]</a> The exact number of genes encoded by the nucleus and the <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial genome</a> differs between species. Most mitochondrial genomes are circular.<a href="#cite_note-Fukuhara-1993-145">[145]</a> In general, mitochondrial DNA lacks <a href="/wiki/Intron" title="Intron">introns</a>, as is the case in the human mitochondrial genome;<a href="#cite_note-Anderson-1981-144">[144]</a> however, introns have been observed in some eukaryotic mitochondrial DNA,<a href="#cite_note-146">[146]</a> such as that of <a href="/wiki/Yeast" title="Yeast">yeast</a><a href="#cite_note-147">[147]</a> and <a href="/wiki/Protist" title="Protist">protists</a>,<a href="#cite_note-148">[148]</a> including <a href="/wiki/Dictyostelium" title="Dictyostelium">Dictyostelium</a> discoideum.<a href="#cite_note-149">[149]</a> Between protein-coding regions, tRNAs are present. Mitochondrial tRNA genes have different sequences from the nuclear tRNAs, but lookalikes of mitochondrial tRNAs have been found in the nuclear chromosomes with high sequence similarity.<a href="#cite_note-Telonis-2014-150">[150]</a> In animals, the mitochondrial genome is typically a single circular chromosome that is approximately 16 kb long and has 37 genes. The genes, while highly conserved, may vary in location. Curiously, this pattern is not found in the human body louse (<a href="/wiki/Pediculus_humanus" title="Pediculus humanus">Pediculus humanus</a>). Instead, this mitochondrial genome is arranged in 18 minicircular chromosomes, each of which is 3–4 kb long and has one to three genes.<a href="#cite_note-Shao-2009-151">[151]</a> This pattern is also found in other <a href="/wiki/Anoplura" class="mw-redirect" title="Anoplura">sucking lice</a>, but not in <a href="/wiki/Mallophaga" title="Mallophaga">chewing lice</a>. Recombination has been shown to occur between the minichromosomes. <div class="mw-heading mw-heading3"><h3 id="Human_population_genetic_studies">Human population genetic studies</h3></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/Human_mitochondrial_genetics" title="Human mitochondrial genetics">Human mitochondrial genetics</a></div> The near-absence of <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> in mitochondrial DNA makes it a useful source of information for studying <a href="/wiki/Population_genetics" title="Population genetics">population genetics</a> and <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>.<a href="#cite_note-152">[152]</a> Because all the mitochondrial DNA is inherited as a single unit, or <a href="/wiki/Haplotype" title="Haplotype">haplotype</a>, the relationships between mitochondrial DNA from different individuals can be represented as a <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">gene tree</a>. Patterns in these gene trees can be used to infer the evolutionary history of populations. The classic example of this is in <a href="/wiki/Human_evolutionary_genetics" title="Human evolutionary genetics">human evolutionary genetics</a>, where the <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a> can be used to provide a recent date for <a href="/wiki/Mitochondrial_Eve" title="Mitochondrial Eve">mitochondrial Eve</a>.<a href="#cite_note-153">[153]</a><a href="#cite_note-154">[154]</a> This is often interpreted as strong support for a recent modern human expansion <a href="/wiki/Recent_single-origin_hypothesis" class="mw-redirect" title="Recent single-origin hypothesis">out of Africa</a>.<a href="#cite_note-Garrigan-2006-155">[155]</a> Another human example is the sequencing of mitochondrial DNA from <a href="/wiki/Neanderthal" title="Neanderthal">Neanderthal</a> bones. The relatively large evolutionary distance between the mitochondrial DNA sequences of Neanderthals and living humans has been interpreted as evidence for the lack of interbreeding between Neanderthals and modern humans.<a href="#cite_note-156">[156]</a> However, mitochondrial DNA reflects only the history of the females in a population. This can be partially overcome by the use of paternal genetic sequences, such as the <a href="/wiki/Genetic_recombination" title="Genetic recombination">non-recombining</a> region of the <a href="/wiki/Y-chromosome" class="mw-redirect" title="Y-chromosome">Y-chromosome</a>.<a href="#cite_note-Garrigan-2006-155">[155]</a> Recent measurements of the <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a> for mitochondrial DNA<a href="#cite_note-157">[157]</a> reported a value of 1 mutation every 7884 years dating back to the most recent common ancestor of humans and apes, which is consistent with estimates of mutation rates of autosomal DNA (10−8 per base per generation).<a href="#cite_note-Nachman-2000-158">[158]</a> <div class="mw-heading mw-heading3"><h3 id="Alternative_genetic_code">Alternative genetic code</h3></div> <div style="border: 1px solid grey; float:right; margin-right:0.6em; padding:0.5em"> <table class="wikitable"> <caption>Exceptions to the standard genetic code in mitochondria<a href="#cite_note-Alberts-2005-20">[20]</a> </caption> <tbody><tr> <th>Organism</th> <th>Codon</th> <th>Standard</th> <th>Mitochondria </th></tr> <tr> <td><a href="/wiki/Mammal" title="Mammal">Mammals</a> </td> <td>AGA, AGG </td> <td>Arginine </td> <td>Stop codon </td></tr> <tr> <td><a href="/wiki/Invertebrate" title="Invertebrate">Invertebrates</a> </td> <td>AGA, AGG </td> <td>Arginine </td> <td>Serine </td></tr> <tr> <td><a href="/wiki/Fungus" title="Fungus">Fungi</a> </td> <td>CUA </td> <td>Leucine </td> <td>Threonine </td></tr> <tr valign="top"> <td rowspan="2">All of the above </td> <td>AUA </td> <td>Isoleucine </td> <td>Methionine </td></tr> <tr> <td>UGA </td> <td>Stop codon </td> <td>Tryptophan </td></tr></tbody></table> </div> While slight variations on the standard <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> had been predicted earlier,<a href="#cite_note-159">[159]</a> none was discovered until 1979, when researchers studying <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">human mitochondrial genes</a> determined that they used an alternative code.<a href="#cite_note-160">[160]</a> Nonetheless, the mitochondria of many other eukaryotes, including most plants, use the standard code.<a href="#cite_note-Elzanowski-2019-161">[161]</a> Many slight variants have been discovered since,<a href="#cite_note-Elzanowski-2019-161">[161]</a> including various alternative mitochondrial codes.<a href="#cite_note-162">[162]</a> Further, the AUA, AUC, and AUU codons are all allowable start codons. Some of these differences should be regarded as pseudo-changes in the genetic code due to the phenomenon of <a href="/wiki/RNA_editing" title="RNA editing">RNA editing</a>, which is common in mitochondria. In higher plants, it was thought that CGG encoded for <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> and not <a href="/wiki/Arginine" title="Arginine">arginine</a>; however, the codon in the processed RNA was discovered to be the UGG codon, consistent with the standard <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> for tryptophan.<a href="#cite_note-163">[163]</a> Of note, the arthropod mitochondrial genetic code has undergone parallel evolution within a phylum, with some organisms uniquely translating AGG to lysine.<a href="#cite_note-164">[164]</a> <div class="mw-heading mw-heading3"><h3 id="Replication_and_inheritance">Replication and inheritance</h3></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/Mitochondrial_fission" title="Mitochondrial fission">Mitochondrial fission</a></div> Mitochondria divide by <a href="/wiki/Mitochondrial_fission" title="Mitochondrial fission">mitochondrial fission</a>, a form of <a href="/wiki/Binary_fission" class="mw-redirect" title="Binary fission">binary fission</a> that is also done by bacteria<a href="#cite_note-165">[165]</a> although the process is tightly regulated by the host eukaryotic cell and involves communication between and contact with several other organelles. The regulation of this division differs between eukaryotes. In many single-celled eukaryotes, their growth and division are linked to the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>. For example, a single mitochondrion may divide synchronously with the nucleus. This division and segregation process must be tightly controlled so that each daughter cell receives at least one mitochondrion. In other eukaryotes (in mammals for example), mitochondria may replicate their DNA and divide mainly in response to the energy needs of the cell, rather than in phase with the cell cycle. When the energy needs of a cell are high, mitochondria grow and divide. When energy use is low, mitochondria are destroyed or become inactive. In such examples mitochondria are apparently randomly distributed to the daughter cells during the division of the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. Mitochondrial dynamics, the balance between <a href="/wiki/Mitochondrial_fusion" title="Mitochondrial fusion">mitochondrial fusion</a> and <a href="/wiki/Mitochondrial_fission" title="Mitochondrial fission">fission</a>, is an important factor in pathologies associated with several disease conditions.<a href="#cite_note-166">[166]</a> The hypothesis of mitochondrial binary fission has relied on the visualization by fluorescence microscopy and conventional <a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">transmission electron microscopy</a> (TEM). The resolution of fluorescence microscopy (≈200 nm) is insufficient to distinguish structural details, such as double mitochondrial membrane in mitochondrial division or even to distinguish individual mitochondria when several are close together. Conventional TEM has also some technical limitations[<a href="/wiki/Wikipedia:Avoid_weasel_words" class="mw-redirect" title="Wikipedia:Avoid weasel words">which?</a>] in verifying mitochondrial division. <a href="/wiki/Cryo-electron_tomography" class="mw-redirect" title="Cryo-electron tomography">Cryo-electron tomography</a> was recently used to visualize mitochondrial division in frozen hydrated intact cells. It revealed that mitochondria divide by budding.<a href="#cite_note-167">[167]</a> An individual's mitochondrial genes are inherited only from the mother, with rare exceptions.<a href="#cite_note-McWilliams-2019-168">[168]</a> In humans, when an <a href="/wiki/Ovum" class="mw-redirect" title="Ovum">egg cell</a> is fertilized by a sperm, the mitochondria, and therefore the mitochondrial DNA, usually come from the egg only. The sperm's mitochondria enter the egg, but do not contribute genetic information to the embryo.<a href="#cite_note-169">[169]</a> Instead, paternal mitochondria are marked with <a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitin</a> to select them for later destruction inside the <a href="/wiki/Embryo" title="Embryo">embryo</a>.<a href="#cite_note-170">[170]</a> The egg cell contains relatively few mitochondria, but these mitochondria divide to populate the cells of the adult organism. This mode is seen in most organisms, including the majority of animals. However, mitochondria in some species can sometimes be inherited paternally. This is the norm among certain <a href="/wiki/Conifer" title="Conifer">coniferous</a> plants, although not in <a href="/wiki/Pine_tree" class="mw-redirect" title="Pine tree">pine trees</a> and <a href="/wiki/Taxus" title="Taxus">yews</a>.<a href="#cite_note-171">[171]</a> For <a href="/wiki/Mytilidae" title="Mytilidae">Mytilids</a>, paternal inheritance only occurs within males of the species.<a href="#cite_note-172">[172]</a><a href="#cite_note-173">[173]</a><a href="#cite_note-174">[174]</a> It has been suggested that it occurs at a very low level in humans.<a href="#cite_note-175">[175]</a> <a href="/wiki/Uniparental_inheritance" title="Uniparental inheritance">Uniparental inheritance</a> leads to little opportunity for <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> between different lineages of mitochondria, although a single mitochondrion can contain 2–10 copies of its DNA.<a href="#cite_note-Wiesner-1992-142">[142]</a> What recombination does take place maintains genetic integrity rather than maintaining diversity. However, there are studies showing evidence of recombination in mitochondrial DNA. It is clear that the enzymes necessary for recombination are present in mammalian cells.<a href="#cite_note-176">[176]</a> Further, evidence suggests that animal mitochondria can undergo recombination.<a href="#cite_note-177">[177]</a> The data are more controversial in humans, although indirect evidence of recombination exists.<a href="#cite_note-178">[178]</a><a href="#cite_note-179">[179]</a> Entities undergoing uniparental inheritance and with little to no recombination may be expected to be subject to <a href="/wiki/Muller%27s_ratchet" title="Muller's ratchet">Muller's ratchet</a>, the accumulation of deleterious mutations until functionality is lost. Animal populations of mitochondria avoid this buildup through a developmental process known as the <a href="/wiki/Heteroplasmy#Mitochondrial_bottleneck" title="Heteroplasmy">mtDNA bottleneck</a>. The bottleneck exploits <a href="/wiki/Cellular_noise" title="Cellular noise">stochastic processes in the cell</a> to increase the cell-to-cell variability in <a href="/wiki/Heteroplasmy" title="Heteroplasmy">mutant load</a> as an organism develops: a single egg cell with some proportion of mutant mtDNA thus produces an embryo where different cells have different mutant loads. Cell-level selection may then act to remove those cells with more mutant mtDNA, leading to a stabilization or reduction in mutant load between generations. The mechanism underlying the bottleneck is debated,<a href="#cite_note-180">[180]</a><a href="#cite_note-181">[181]</a><a href="#cite_note-182">[182]</a> with a recent mathematical and experimental metastudy providing evidence for a combination of random partitioning of mtDNAs at cell divisions and random turnover of mtDNA molecules within the cell.<a href="#cite_note-183">[183]</a> <div class="mw-heading mw-heading3"><h3 id="DNA_repair">DNA repair</h3></div> Mitochondria can repair oxidative <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage</a> by mechanisms analogous to those occurring in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>. The proteins employed in <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mtDNA</a> repair are encoded by nuclear <a href="/wiki/Gene" title="Gene">genes</a>, and are translocated to the mitochondria. The <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> pathways in mammalian mitochondria include <a href="/wiki/Base_excision_repair" title="Base excision repair">base excision repair</a>, double-strand break repair, direct reversal and <a href="/wiki/DNA_mismatch_repair" title="DNA mismatch repair">mismatch repair</a>.<a href="#cite_note-Gredilla-2012-184">[184]</a><a href="#cite_note-Saki-2017-185">[185]</a> Alternatively, DNA damage may be bypassed, rather than repaired, by translesion synthesis. Of the several DNA repair process in mitochondria, the base excision repair pathway has been most comprehensively studied.<a href="#cite_note-Saki-2017-185">[185]</a> Base excision repair is carried out by a sequence of enzyme-catalyzed steps that include recognition and excision of a damaged DNA base, removal of the resulting abasic site, end processing, gap filling and ligation. A common damage in mtDNA that is repaired by base excision repair is <a href="/wiki/8-oxoguanine" class="mw-redirect" title="8-oxoguanine">8-oxoguanine</a> produced by oxidation of <a href="/wiki/Guanine" title="Guanine">guanine</a>.<a href="#cite_note-Leon-2016-186">[186]</a> Double-strand breaks can be repaired by <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombinational</a> repair in both mammalian mtDNA<a href="#cite_note-Dahal-2018-187">[187]</a> and plant mtDNA.<a href="#cite_note-Odahara-2007-188">[188]</a> Double-strand breaks in mtDNA can also be repaired by <a href="/wiki/Microhomology-mediated_end_joining" title="Microhomology-mediated end joining">microhomology-mediated end joining</a>.<a href="#cite_note-Tadi-2016-189">[189]</a> Although there is evidence for the repair processes of direct reversal and mismatch repair in mtDNA, these processes are not well characterized.<a href="#cite_note-Saki-2017-185">[185]</a> <div class="mw-heading mw-heading3"><h3 id="Lack_of_mitochondrial_DNA">Lack of mitochondrial DNA</h3></div> Some organisms have lost mitochondrial DNA altogether. In these cases, genes encoded by the mitochondrial DNA have been lost or transferred to the nucleus.<a href="#cite_note-Chan-2006-141">[141]</a> <a href="/wiki/Cryptosporidium" title="Cryptosporidium">Cryptosporidium</a> have mitochondria that lack any DNA, presumably because all their genes have been lost or transferred.<a href="#cite_note-Henriquez-2005-190">[190]</a> In Cryptosporidium, the mitochondria have an altered <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> generation system that renders the parasite resistant to many classical mitochondrial <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">inhibitors</a> such as <a href="/wiki/Cyanide" title="Cyanide">cyanide</a>, <a href="/wiki/Azide" title="Azide">azide</a>, and <a href="/wiki/Atovaquone" title="Atovaquone">atovaquone</a>.<a href="#cite_note-Henriquez-2005-190">[190]</a> Mitochondria that lack their own DNA have been found in a marine parasitic <a href="/wiki/Dinoflagellate" title="Dinoflagellate">dinoflagellate</a> from the genus <a href="/wiki/Amoebophyra" title="Amoebophyra">Amoebophyra</a>. This microorganism, A. cerati, has functional mitochondria that lack a genome.<a href="#cite_note-191">[191]</a> In related species, the mitochondrial genome still has three genes, but in A. cerati only a single mitochondrial gene — the <a href="/wiki/Cytochrome_c_oxidase_subunit_I" title="Cytochrome c oxidase subunit I">cytochrome c oxidase I</a> gene (cox1) — is found, and it has migrated to the genome of the nucleus.<a href="#cite_note-192">[192]</a> <div class="mw-heading mw-heading2"><h2 id="Dysfunction_and_disease">Dysfunction and disease</h2></div> <div class="mw-heading mw-heading3"><h3 id="Mitochondrial_diseases">Mitochondrial diseases</h3></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/Mitochondrial_disease" title="Mitochondrial disease">Mitochondrial disease</a></div> Damage and subsequent dysfunction in mitochondria is an important factor in a range of human diseases due to their influence in cell metabolism. Mitochondrial disorders often present as neurological disorders, including <a href="/wiki/Autism" title="Autism">autism</a>.<a href="#cite_note-Griffiths-2017-18">[18]</a> They can also manifest as <a href="/wiki/Myopathy" title="Myopathy">myopathy</a>, <a href="/wiki/Diabetes" title="Diabetes">diabetes</a>, multiple <a href="/wiki/Endocrinopathy" class="mw-redirect" title="Endocrinopathy">endocrinopathy</a>, and a variety of other systemic disorders.<a href="#cite_note-Zeviani-2004-193">[193]</a> Diseases caused by mutation in the mtDNA include <a href="/wiki/Kearns%E2%80%93Sayre_syndrome" title="Kearns–Sayre syndrome">Kearns–Sayre syndrome</a>, <a href="/wiki/MELAS_syndrome" title="MELAS syndrome">MELAS syndrome</a> and <a href="/wiki/Leber%27s_hereditary_optic_neuropathy" title="Leber's hereditary optic neuropathy">Leber's hereditary optic neuropathy</a>.<a href="#cite_note-Taylor-2005-194">[194]</a> In the vast majority of cases, these diseases are transmitted by a female to her children, as the <a href="/wiki/Zygote" title="Zygote">zygote</a> derives its mitochondria and hence its mtDNA from the ovum. Diseases such as Kearns-Sayre syndrome, <a href="/wiki/Pearson_syndrome" title="Pearson syndrome">Pearson syndrome</a>, and progressive external <a href="/wiki/Ophthalmoparesis" title="Ophthalmoparesis">ophthalmoplegia</a> are thought to be due to large-scale mtDNA rearrangements, whereas other diseases such as MELAS syndrome, Leber's hereditary optic neuropathy, <a href="/wiki/MERRF_syndrome" title="MERRF syndrome">MERRF syndrome</a>, and others are due to <a href="/wiki/Point_mutation" title="Point mutation">point mutations</a> in mtDNA.<a href="#cite_note-Zeviani-2004-193">[193]</a> It has also been reported that drug tolerant cancer cells have an increased number and size of mitochondria which suggested an increase in mitochondrial biogenesis.<a href="#cite_note-Goldman-2019-195">[195]</a> A 2022 study in Nature Nanotechnology has reported that cancer cells can hijack the mitochondria from immune cells via physical tunneling nanotubes.<a href="#cite_note-196">[196]</a> In other diseases, defects in nuclear genes lead to dysfunction of mitochondrial proteins. This is the case in <a href="/wiki/Friedreich%27s_ataxia" title="Friedreich's ataxia">Friedreich's ataxia</a>, <a href="/wiki/Hereditary_spastic_paraplegia" title="Hereditary spastic paraplegia">hereditary spastic paraplegia</a>, and <a href="/wiki/Wilson%27s_disease" title="Wilson's disease">Wilson's disease</a>.<a href="#cite_note-197">[197]</a> These diseases are inherited in a <a href="/wiki/Dominance_relationship" class="mw-redirect" title="Dominance relationship">dominance relationship</a>, as applies to most other genetic diseases. A variety of disorders can be caused by nuclear mutations of oxidative phosphorylation enzymes, such as <a href="/wiki/Coenzyme_Q10" title="Coenzyme Q10">coenzyme Q10</a> deficiency and <a href="/wiki/Barth_syndrome" title="Barth syndrome">Barth syndrome</a>.<a href="#cite_note-Zeviani-2004-193">[193]</a> Environmental influences may interact with hereditary predispositions and cause mitochondrial disease. For example, there may be a link between <a href="/wiki/Pesticide" title="Pesticide">pesticide</a> exposure and the later onset of <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a>.<a href="#cite_note-198">[198]</a><a href="#cite_note-199">[199]</a> Other pathologies with etiology involving mitochondrial dysfunction include <a href="/wiki/Schizophrenia" title="Schizophrenia">schizophrenia</a>, <a href="/wiki/Bipolar_disorder" title="Bipolar disorder">bipolar disorder</a>, <a href="/wiki/Dementia" title="Dementia">dementia</a>, <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a>,<a href="#cite_note-200">[200]</a><a href="#cite_note-201">[201]</a> Parkinson's disease, <a href="/wiki/Epilepsy" title="Epilepsy">epilepsy</a>, <a href="/wiki/Stroke" title="Stroke">stroke</a>, <a href="/wiki/Cardiovascular_disease" title="Cardiovascular disease">cardiovascular disease</a>, <a href="/wiki/Myalgic_encephalomyelitis/chronic_fatigue_syndrome" title="Myalgic encephalomyelitis/chronic fatigue syndrome">myalgic encephalomyelitis/chronic fatigue syndrome</a> (ME/CFS), <a href="/wiki/Retinitis_pigmentosa" title="Retinitis pigmentosa">retinitis pigmentosa</a>, and <a href="/wiki/Diabetes_mellitus" class="mw-redirect" title="Diabetes mellitus">diabetes mellitus</a>.<a href="#cite_note-202">[202]</a><a href="#cite_note-Pieczenik-2007-203">[203]</a> Mitochondria-mediated oxidative stress plays a role in cardiomyopathy in <a href="/wiki/Type_2_diabetics" class="mw-redirect" title="Type 2 diabetics">type 2 diabetics</a>. Increased fatty acid delivery to the heart increases fatty acid uptake by cardiomyocytes, resulting in increased fatty acid oxidation in these cells. This process increases the reducing equivalents available to the electron transport chain of the mitochondria, ultimately increasing reactive oxygen species (ROS) production. ROS increases <a href="/wiki/Uncoupling_proteins" class="mw-redirect" title="Uncoupling proteins">uncoupling proteins</a> (UCPs) and potentiate proton leakage through the <a href="/wiki/Adenine_nucleotide_translocator" title="Adenine nucleotide translocator">adenine nucleotide translocator</a> (ANT), the combination of which <a href="/wiki/Uncoupler" title="Uncoupler">uncouples</a> the mitochondria. Uncoupling then increases oxygen consumption by the mitochondria, compounding the increase in fatty acid oxidation. This creates a vicious cycle of uncoupling; furthermore, even though oxygen consumption increases, ATP synthesis does not increase proportionally because the mitochondria are uncoupled. Less ATP availability ultimately results in an energy deficit presenting as reduced cardiac efficiency and contractile dysfunction. To compound the problem, impaired sarcoplasmic reticulum calcium release and reduced mitochondrial reuptake limits peak cytosolic levels of the important signaling ion during muscle contraction. Decreased intra-mitochondrial calcium concentration increases dehydrogenase activation and ATP synthesis. So in addition to lower ATP synthesis due to fatty acid oxidation, ATP synthesis is impaired by poor calcium signaling as well, causing cardiac problems for diabetics.<a href="#cite_note-204">[204]</a> Mitochondria also modulate processes such as testicular somatic cell development, spermatogonial stem cell differentiation, luminal acidification, testosterone production in testes, and more. Thus, dysfunction of mitochondria in spermatozoa can be a cause for infertility.<a href="#cite_note-205">[205]</a> In efforts to combat mitochondrial disease, <a href="/wiki/Mitochondrial_replacement_therapy" title="Mitochondrial replacement therapy">mitochondrial replacement therapy</a> (MRT) has been developed. This form of in vitro fertilization uses donor mitochondria, which avoids the transmission of diseases caused by mutations of mitochondrial DNA.<a href="#cite_note-206">[206]</a> However, this therapy is still being researched and can introduce genetic modification, as well as safety concerns. These diseases are rare but can be extremely debilitating and progressive diseases, thus posing complex ethical questions for public policy.<a href="#cite_note-207">[207]</a> <div class="mw-heading mw-heading3"><h3 id="Relationships_to_aging">Relationships to aging</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Hallmarks_of_aging#Mitochondrial_dysfunction" title="Hallmarks of aging">Hallmarks of aging § Mitochondrial dysfunction</a></div> There may be some leakage of the <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electrons transferred</a> in the respiratory chain to form <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a>. This was thought to result in significant <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a> in the mitochondria with high mutation rates of mitochondrial DNA.<a href="#cite_note-208">[208]</a> Hypothesized links between aging and oxidative stress are not new and were proposed in 1956,<a href="#cite_note-209">[209]</a> which was later refined into the <a href="/wiki/Mitochondrial_theory_of_ageing" title="Mitochondrial theory of ageing">mitochondrial free radical theory of aging</a>.<a href="#cite_note-210">[210]</a> A vicious cycle was thought to occur, as oxidative stress leads to mitochondrial DNA mutations, which can lead to enzymatic abnormalities and further oxidative stress. A number of changes can occur to mitochondria during the aging process.<a href="#cite_note-211">[211]</a> Tissues from elderly humans show a decrease in enzymatic activity of the proteins of the respiratory chain.<a href="#cite_note-212">[212]</a> However, mutated mtDNA can only be found in about 0.2% of very old cells.<a href="#cite_note-213">[213]</a> Large deletions in the mitochondrial genome have been hypothesized to lead to high levels of <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a> and neuronal death in <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a>.<a href="#cite_note-214">[214]</a> Mitochondrial dysfunction has also been shown to occur in <a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">amyotrophic lateral sclerosis</a>.<a href="#cite_note-215">[215]</a><a href="#cite_note-216">[216]</a> Since mitochondria cover a pivotal role in the ovarian function, by providing ATP necessary for the development from germinal vesicle to mature <a href="/wiki/Oocyte" title="Oocyte">oocyte</a>, a decreased mitochondria function can lead to inflammation, resulting in premature ovarian failure and accelerated ovarian aging. The resulting dysfunction is then reflected in quantitative (such as mtDNA copy number and mtDNA deletions), qualitative (such as mutations and strand breaks) and oxidative damage (such as dysfunctional mitochondria due to ROS), which are not only relevant in ovarian aging, but perturb oocyte-cumulus crosstalk in the ovary, are linked to genetic disorders (such as Fragile X) and can interfere with embryo selection.<a href="#cite_note-217">[217]</a> <div class="mw-heading mw-heading2"><h2 id="History">History</h2></div> The first observations of intracellular structures that probably represented mitochondria were published in 1857, by the physiologist <a href="/wiki/Albert_von_K%C3%B6lliker" title="Albert von Kölliker">Albert von Kolliker</a>.<a href="#cite_note-218">[218]</a><a href="#cite_note-Ernster-1981-219">[219]</a> <a href="/wiki/Richard_Altmann" title="Richard Altmann">Richard Altmann</a>, in 1890, established them as cell organelles and called them "bioblasts".<a href="#cite_note-Ernster-1981-219">[219]</a><a href="#cite_note-220">[220]</a> In 1898, Carl Benda coined the term "mitochondria" from the <a href="/wiki/Greek_language" title="Greek language">Greek</a> μίτος, mitos, "thread", and χονδρίον, chondrion, "granule".<a href="#cite_note-OnlineEtymDict-221">[221]</a><a href="#cite_note-Ernster-1981-219">[219]</a><a href="#cite_note-222">[222]</a> <a href="/wiki/Leonor_Michaelis" title="Leonor Michaelis">Leonor Michaelis</a> discovered that <a href="/wiki/Janus_green" class="mw-redirect" title="Janus green">Janus green</a> can be used as a <a href="/wiki/Supravital_stain" class="mw-redirect" title="Supravital stain">supravital stain</a> for mitochondria in 1900.<a href="#cite_note-223">[223]</a> In 1904, <a href="/w/index.php?title=Friedrich_Meves&action=edit&redlink=1" class="new" title="Friedrich Meves (page does not exist)">Friedrich Meves</a> made the first recorded observation of mitochondria in plants in cells of the white waterlily, <a href="/wiki/Nymphaea_alba" title="Nymphaea alba">Nymphaea alba</a>,<a href="#cite_note-Ernster-1981-219">[219]</a><a href="#cite_note-224">[224]</a> and in 1908, along with <a href="/wiki/Claudius_Regaud" title="Claudius Regaud">Claudius Regaud</a>, suggested that they contain proteins and lipids. Benjamin F. Kingsbury, in 1912, first related them with cell respiration, but almost exclusively based on morphological observations.<a href="#cite_note-225">[225]</a><a href="#cite_note-Ernster-1981-219">[219]</a> In 1913, <a href="/wiki/Otto_Heinrich_Warburg" title="Otto Heinrich Warburg">Otto Heinrich Warburg</a> linked respiration to particles which he had obtained from extracts of guinea-pig liver and which he called "grana".<a href="#cite_note-226">[226]</a> Warburg and <a href="/wiki/Heinrich_Otto_Wieland" title="Heinrich Otto Wieland">Heinrich Otto Wieland</a>, who had also postulated a similar particle mechanism, disagreed on the chemical nature of the respiration. It was not until 1925, when <a href="/wiki/David_Keilin" title="David Keilin">David Keilin</a> discovered <a href="/wiki/Cytochromes" class="mw-redirect" title="Cytochromes">cytochromes</a>, that the <a href="/wiki/Respiratory_chain" class="mw-redirect" title="Respiratory chain">respiratory chain</a> was described.<a href="#cite_note-Ernster-1981-219">[219]</a> In 1939, experiments using minced muscle cells demonstrated that cellular respiration using one <a href="/wiki/Oxygen" title="Oxygen">oxygen molecule</a> can form four <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP) molecules, and in 1941, the concept of the phosphate bonds of ATP being a form of energy in cellular metabolism was developed by <a href="/wiki/Fritz_Albert_Lipmann" title="Fritz Albert Lipmann">Fritz Albert Lipmann</a>. In the following years, the mechanism behind cellular respiration was further elaborated, although its link to the mitochondria was not known.<a href="#cite_note-Ernster-1981-219">[219]</a> The introduction of <a href="/wiki/Cell_fractionation" title="Cell fractionation">tissue fractionation</a> by <a href="/wiki/Albert_Claude" title="Albert Claude">Albert Claude</a> allowed mitochondria to be isolated from other cell fractions and biochemical analysis to be conducted on them alone. In 1946, he concluded that <a href="/wiki/Cytochrome_oxidase" class="mw-redirect" title="Cytochrome oxidase">cytochrome oxidase</a> and other enzymes responsible for the respiratory chain were isolated to the mitochondria. <a href="/wiki/Eugene_P._Kennedy" title="Eugene P. Kennedy">Eugene Kennedy</a> and <a href="/wiki/Albert_L._Lehninger" title="Albert L. Lehninger">Albert Lehninger</a> discovered in 1948 that mitochondria are the site of <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a> in eukaryotes. Over time, the fractionation method was further developed, improving the quality of the mitochondria isolated, and other elements of <a href="/wiki/Cellular_respiration" title="Cellular respiration">cell respiration</a> were determined to occur in the mitochondria.<a href="#cite_note-Ernster-1981-219">[219]</a> The first high-resolution electron <a href="/wiki/Micrographs" class="mw-redirect" title="Micrographs">micrographs</a> appeared in 1952, replacing the Janus Green stains as the preferred way to visualize mitochondria.<a href="#cite_note-Ernster-1981-219">[219]</a> This led to a more detailed analysis of the structure of the mitochondria, including confirmation that they were surrounded by a membrane. It also showed a second membrane inside the mitochondria that folded up in ridges dividing up the inner chamber and that the size and shape of the mitochondria varied from cell to cell. The popular term "powerhouse of the cell" was coined by <a href="/wiki/Philip_Siekevitz" title="Philip Siekevitz">Philip Siekevitz</a> in 1957.<a href="#cite_note-Siekevitz-1957-4">[4]</a><a href="#cite_note-227">[227]</a> In 1967, it was discovered that mitochondria contained <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a>.<a href="#cite_note-228">[228]</a> In 1968, methods were developed for mapping the mitochondrial genes, with the genetic and physical map of yeast mitochondrial DNA completed in 1976.<a href="#cite_note-Ernster-1981-219">[219]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div> <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: 20em;"> <ul><li><a href="/wiki/Anti-mitochondrial_antibodies" class="mw-redirect" title="Anti-mitochondrial antibodies">Anti-mitochondrial antibodies</a></li> <li><a href="/wiki/Respirometry#Mitochondrial_metabolic_rates" title="Respirometry">Mitochondrial metabolic rates</a></li> <li><a href="/wiki/Mitochondrial_permeability_transition_pore" title="Mitochondrial permeability transition pore">Mitochondrial permeability transition pore</a></li> <li><a href="/wiki/Mitophagy" title="Mitophagy">Mitophagy</a></li> <li><a href="/wiki/Nebenkern" title="Nebenkern">Nebenkern</a></li> <li><a href="/wiki/Oncocyte" title="Oncocyte">Oncocyte</a></li> <li><a href="/wiki/Oncocytoma" title="Oncocytoma">Oncocytoma</a></li> <li><a href="/wiki/Paternal_mtDNA_transmission" title="Paternal mtDNA transmission">Paternal mtDNA transmission</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li> <li><a href="/wiki/Submitochondrial_particle" title="Submitochondrial particle">Submitochondrial particle</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 32em;"> <ol class="references"> <li id="cite_note-1"><a href="#cite_ref-1">^</a> <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 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Acta Neuropathologica. 141 (2): 257–279. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00401-020-02252-5">10.1007/s00401-020-02252-5</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7847443">7847443</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/33398403">33398403</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Acta+Neuropathologica&rft.atitle=Mitochondrial+bioenergetic+deficits+in+C9orf72+amyotrophic+lateral+sclerosis+motor+neurons+cause+dysfunctional+axonal+homeostasis&rft.volume=141&rft.issue=2&rft.pages=257-279&rft.date=2021-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7847443%23id-name%3DPMC&rft_id=info%3Apmid%2F33398403&rft_id=info%3Adoi%2F10.1007%2Fs00401-020-02252-5&rft.aulast=Mehta&rft.aufirst=AR&rft.au=Gregory%2C+JM&rft.au=Dando%2C+O&rft.au=Carter%2C+RN&rft.au=Burr%2C+K&rft.au=Nanda%2C+J&rft.au=Story%2C+D&rft.au=McDade%2C+K&rft.au=Smith%2C+C&rft.au=Morton%2C+NM&rft.au=Mahad%2C+DJ&rft.au=Hardingham%2C+GE&rft.au=Chandran%2C+S&rft.au=Selvaraj%2C+BT&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7847443&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-217"><a href="#cite_ref-217">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChiangShuklaPagidasAhmed2020" class="citation journal cs1">Chiang JL, Shukla P, Pagidas K, Ahmed NS, Karri S, Gunn DD, et al. (November 2020). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9375691">"Mitochondria in Ovarian Aging and Reproductive Longevity"</a>. Ageing Research Reviews. 63: 101168. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.arr.2020.101168">10.1016/j.arr.2020.101168</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9375691">9375691</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/32896666">32896666</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:221472721">221472721</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ageing+Research+Reviews&rft.atitle=Mitochondria+in+Ovarian+Aging+and+Reproductive+Longevity&rft.volume=63&rft.pages=101168&rft.date=2020-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC9375691%23id-name%3DPMC&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A221472721%23id-name%3DS2CID&rft_id=info%3Apmid%2F32896666&rft_id=info%3Adoi%2F10.1016%2Fj.arr.2020.101168&rft.aulast=Chiang&rft.aufirst=JL&rft.au=Shukla%2C+P&rft.au=Pagidas%2C+K&rft.au=Ahmed%2C+NS&rft.au=Karri%2C+S&rft.au=Gunn%2C+DD&rft.au=Hurd%2C+WW&rft.au=Singh%2C+KK&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC9375691&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-218"><a href="#cite_ref-218">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKölliker1857" class="citation journal cs1 cs1-prop-foreign-lang-source">Kölliker A (1857). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/49190#page/319/mode/1up">"Einige Bemerkungen über die Endigungen der Hautnerven und den Bau der Muskeln"</a> [Some remarks about the terminations of the cutaneous nerves and the structure of muscles]. Zeitschrift für wissenschaftliche Zoologie (in German). 8: 311–325. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230622033955/https://www.biodiversitylibrary.org/item/49190#page/319/mode/1up">Archived</a> from the original on June 22, 2023. Retrieved June 22, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+wissenschaftliche+Zoologie&rft.atitle=Einige+Bemerkungen+%C3%BCber+die+Endigungen+der+Hautnerven+und+den+Bau+der+Muskeln&rft.volume=8&rft.pages=311-325&rft.date=1857&rft.aulast=K%C3%B6lliker&rft.aufirst=A&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F49190%23page%2F319%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> On p. 316, Kölliker described mitochondria which he observed in fresh frog muscles: " ... sehr blasse rundliche Körnchen, welche in langen linienförmigen Zügen [...] wenn man einmal auf dieselben aufmerksam geworden ist." ( ... [they are] very faint round granules, which are embedded in the [muscle's] contractile substance in long linear trains. These granules are located in the whole thickness of the muscle fiber, on the surface as in the interior, and [they] are so numerous that they appear as a not unimportant element of the muscle fibers, once one has become alert to them.) Kölliker said (p. 321) that he had found mitochondria in the muscles of other animals. In Figure 3 of <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/49190#page/564/mode/1up">Table XIV</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230622033955/https://www.biodiversitylibrary.org/item/49190#page/564/mode/1up">Archived</a> June 22, 2023, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a>, Kölliker depicted mitochondria in frog muscles. </li> <li id="cite_note-Ernster-1981-219">^ <a href="#cite_ref-Ernster-1981_219-0">a</a> <a href="#cite_ref-Ernster-1981_219-1">b</a> <a href="#cite_ref-Ernster-1981_219-2">c</a> <a href="#cite_ref-Ernster-1981_219-3">d</a> <a href="#cite_ref-Ernster-1981_219-4">e</a> <a href="#cite_ref-Ernster-1981_219-5">f</a> <a href="#cite_ref-Ernster-1981_219-6">g</a> <a href="#cite_ref-Ernster-1981_219-7">h</a> <a href="#cite_ref-Ernster-1981_219-8">i</a> <a href="#cite_ref-Ernster-1981_219-9">j</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFErnsterSchatz1981" class="citation journal cs1">Ernster L, Schatz G (December 1981). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2112799">"Mitochondria: a historical review"</a>. 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Leipzig, Germany: Veit & Co. p. 125. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230623002552/https://www.deutschestextarchiv.de/book/view/altmann_elementarorganismen_1890?p=141">Archived</a> from the original on June 23, 2023. Retrieved June 23, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Die+Elementarorganismen+und+ihre+Beziehungen+zu+den+Zellen&rft.place=Leipzig%2C+Germany&rft.pages=125&rft.pub=Veit+%26+Co.&rft.date=1890&rft.aulast=Altmann&rft.aufirst=R&rft_id=https%3A%2F%2Fwww.deutschestextarchiv.de%2Fbook%2Fview%2Faltmann_elementarorganismen_1890%3Fp%3D141&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> From p. 125: "Da auch sonst mancherlei Umstände dafür sprechen, dass Mikroorganismen und Granula einander gleichwerthig sind und Elementarorganismen vorstellen, welche sich überall finden, wo lebendige Kräfte ausgelöst werden, so wollen wir sie mit dem gemeinschaftlichen Namen der Bioblasten bezeichnen." (Since otherwise some circumstances indicate that microorganisms and granula are equivalent to each other and suggest elementary organisms, which are to be found wherever living forces are unleashed, we will designate them with the collective name of "bioblasts".) </li> <li id="cite_note-OnlineEtymDict-221"><a href="#cite_ref-OnlineEtymDict_221-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation encyclopaedia cs1"><a rel="nofollow" class="external text" href="http://www.etymonline.com/index.php?term=mitochondria&allowed_in_frame=0">"mitochondria"</a>. <a href="/wiki/Online_Etymology_Dictionary" title="Online Etymology Dictionary">Online Etymology Dictionary</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160304105458/http://www.etymonline.com/index.php?term=mitochondria&allowed_in_frame=0">Archived</a> from the original on March 4, 2016. Retrieved May 23, 2013.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=mitochondria&rft.btitle=Online+Etymology+Dictionary&rft_id=http%3A%2F%2Fwww.etymonline.com%2Findex.php%3Fterm%3Dmitochondria%26allowed_in_frame%3D0&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-222"><a href="#cite_ref-222">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBenda1898" class="citation journal cs1 cs1-prop-foreign-lang-source">Benda C (1898). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/109725#page/403/mode/1up">"Ueber die Spermatogenese der Vertebraten und höherer Evertebraten. II. Theil: Die Histiogenese der Spermien"</a> [On spermatogenesis in vertebrates and higher invertebrates. Part II: The histogenesis of sperm.]. Archiv für Physiologie (in German). 1898: 393–398. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20190224231523/https://www.biodiversitylibrary.org/item/109725#page/403/mode/1up">Archived</a> from the original on February 24, 2019. Retrieved January 14, 2018.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archiv+f%C3%BCr+Physiologie&rft.atitle=Ueber+die+Spermatogenese+der+Vertebraten+und+h%C3%B6herer+Evertebraten.+II.+Theil%3A+Die+Histiogenese+der+Spermien.&rft.volume=1898&rft.pages=393-398&rft.date=1898&rft.aulast=Benda&rft.aufirst=C&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F109725%23page%2F403%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> From p. 397: After Brenda states that " ... ich bereits in vielen Zellarten aller möglichen Thierclassen gefunden habe, ... " ( ... I have already found [them (mitochondria)] in many types of cells of all possible classes of animals, ... ), he suggests: "Ich möchte vorläufig vorschlagen, ihnen als Mitochondria eine besondere Stellung vorzubehalten, die ich in weiteren Arbeiten begründen werde." (I would like to suggest provisionally reserving for them, as "mitochondria", a special status which I will justify in further work.) <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBenda1899" class="citation journal cs1 cs1-prop-foreign-lang-source">Benda C (1899). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/109704#page/388/mode/1up">"Weitere Mitteilungen über die Mitochondria"</a> [Further reports on mitochondria]. Archiv für Physiologie: Verhandlungen der Berliner Physiologischen Gesellschaft (in German). 1899: 376–383. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230623035548/https://www.biodiversitylibrary.org/item/109704#page/388/mode/1up">Archived</a> from the original on June 23, 2023. Retrieved June 23, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archiv+f%C3%BCr+Physiologie%3A+Verhandlungen+der+Berliner+Physiologischen+Gesellschaft&rft.atitle=Weitere+Mitteilungen+%C3%BCber+die+Mitochondria&rft.volume=1899&rft.pages=376-383&rft.date=1899&rft.aulast=Benda&rft.aufirst=C&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F109704%23page%2F388%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"></li></ul> </li> <li id="cite_note-223"><a href="#cite_ref-223">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichaelis1900" class="citation journal cs1 cs1-prop-foreign-lang-source">Michaelis L (1900). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/47675#page/580/mode/1up">"Die vitale Farbung, eine Darstellungsmethode der Zellgranula"</a> [The vital stain: a method of imaging cell granules]. Archiv für Mikroskopische Anatomie und Entwicklungsgeschichte [Archive for Microscopic Anatomy and Ontogenesis] (in German). 55: 558–575. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF02977747">10.1007/BF02977747</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:85010030">85010030</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230623050932/https://www.biodiversitylibrary.org/item/47675#page/580/mode/1up">Archived</a> from the original on June 23, 2023. Retrieved June 23, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archiv+f%C3%BCr+Mikroskopische+Anatomie+und+Entwicklungsgeschichte&rft.atitle=Die+vitale+Farbung%2C+eine+Darstellungsmethode+der+Zellgranula&rft.volume=55&rft.pages=558-575&rft.date=1900&rft_id=info%3Adoi%2F10.1007%2FBF02977747&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85010030%23id-name%3DS2CID&rft.aulast=Michaelis&rft.aufirst=L&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F47675%23page%2F580%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-224"><a href="#cite_ref-224">^</a> Ernster's citation <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMeves1908" class="citation journal cs1">Meves F (May 1908). <a rel="nofollow" class="external text" href="https://zenodo.org/record/2034475">"Die Chondriosomen als Träger erblicher Anlagen. Cytologische Studien am Hühnerembryo"</a>. Archiv für Mikroskopische Anatomie. 72 (1): 816–867. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF02982402">10.1007/BF02982402</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:85047330">85047330</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200109171106/https://zenodo.org/record/2034475">Archived</a> from the original on January 9, 2020. Retrieved June 24, 2019.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archiv+f%C3%BCr+Mikroskopische+Anatomie&rft.atitle=Die+Chondriosomen+als+Tr%C3%A4ger+erblicher+Anlagen.+Cytologische+Studien+am+H%C3%BChnerembryo&rft.volume=72&rft.issue=1&rft.pages=816-867&rft.date=1908-05&rft_id=info%3Adoi%2F10.1007%2FBF02982402&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85047330%23id-name%3DS2CID&rft.aulast=Meves&rft.aufirst=F&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F2034475&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> is wrong, correct citation is <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMeves1904" class="citation journal cs1">Meves F (1904). "Über das Vorkommen von Mitochondrien bezw. Chondromiten in Pflanzenzellen". Ber. Dtsch. Bot. Ges. 22: 284–286. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1438-8677.1904.tb05237.x">10.1111/j.1438-8677.1904.tb05237.x</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:257822147">257822147</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ber.+Dtsch.+Bot.+Ges.&rft.atitle=%C3%9Cber+das+Vorkommen+von+Mitochondrien+bezw.+Chondromiten+in+Pflanzenzellen&rft.volume=22&rft.pages=284-286&rft.date=1904&rft_id=info%3Adoi%2F10.1111%2Fj.1438-8677.1904.tb05237.x&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A257822147%23id-name%3DS2CID&rft.aulast=Meves&rft.aufirst=F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988">, cited in Meves' 1908 paper and in <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchmidt1913" class="citation journal cs1">Schmidt EW (1913). <a rel="nofollow" class="external text" href="https://archive.org/stream/progressusreibot04lots/progressusreibot04lots_djvu.txt">"Pflanzliche Mitochondrien"</a>. Progressus Rei Botanicae. 4: 164–183. Retrieved September 21, 2012.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Progressus+Rei+Botanicae&rft.atitle=Pflanzliche+Mitochondrien&rft.volume=4&rft.pages=164-183&rft.date=1913&rft.aulast=Schmidt&rft.aufirst=EW&rft_id=https%3A%2F%2Farchive.org%2Fstream%2Fprogressusreibot04lots%2Fprogressusreibot04lots_djvu.txt&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988">, with confirmation of Nymphaea alba </li> <li id="cite_note-225"><a href="#cite_ref-225">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKingsbury1912" class="citation journal cs1">Kingsbury BF (1912). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/87070#page/51/mode/1up">"Cytoplasmic fixation"</a>. The Anatomical Record. 6 (2): 39–52. <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%2Far.1090060202">10.1002/ar.1090060202</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:221400422">221400422</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230623061252/https://www.biodiversitylibrary.org/item/87070#page/51/mode/1up">Archived</a> from the original on June 23, 2023. Retrieved June 23, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Anatomical+Record&rft.atitle=Cytoplasmic+fixation&rft.volume=6&rft.issue=2&rft.pages=39-52&rft.date=1912&rft_id=info%3Adoi%2F10.1002%2Far.1090060202&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A221400422%23id-name%3DS2CID&rft.aulast=Kingsbury&rft.aufirst=BF&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F87070%23page%2F51%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> From p. 47: " ... the mitochondria are the structural expression thereof [i.e., of the chemical reducing processes in the cytoplasm], ... " </li> <li id="cite_note-226"><a href="#cite_ref-226">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWarburg1913" class="citation journal cs1 cs1-prop-foreign-lang-source">Warburg O (1913). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/108058#page/613/mode/1up">"Über sauerstoffatmende Körnchen aus Leberzellen und über Sauerstoffatmung in Berkefeld-Filtraten wässriger Leberextrake"</a> [On respiring granules from liver cells and on respiration in Berkefeld filtrates of aqueous liver extracts]. Pflügers Archiv für die gesamte Physiologie des Menschen und der Tiere (Pfluger's Archive for All Physiology of Humans and Animals) (in German). 154: 599–617. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230623190650/https://www.biodiversitylibrary.org/item/108058#page/613/mode/1up">Archived</a> from the original on June 23, 2023. Retrieved June 23, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Pfl%C3%BCgers+Archiv+f%C3%BCr+die+gesamte+Physiologie+des+Menschen+und+der+Tiere+%28Pfluger%27s+Archive+for+All+Physiology+of+Humans+and+Animals%29&rft.atitle=%C3%9Cber+sauerstoffatmende+K%C3%B6rnchen+aus+Leberzellen+und+%C3%BCber+Sauerstoffatmung+in+Berkefeld-Filtraten+w%C3%A4ssriger+Leberextrake&rft.volume=154&rft.pages=599-617&rft.date=1913&rft.aulast=Warburg&rft.aufirst=O&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F108058%23page%2F613%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-227"><a href="#cite_ref-227">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMilaneTrivediSinghTalekar2015" class="citation journal cs1">Milane L, Trivedi M, Singh A, Talekar M, Amiji M (June 2015). "Mitochondrial biology, targets, and drug delivery". Journal of Controlled Release. 207: 40–58. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jconrel.2015.03.036">10.1016/j.jconrel.2015.03.036</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/25841699">25841699</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+Controlled+Release&rft.atitle=Mitochondrial+biology%2C+targets%2C+and+drug+delivery&rft.volume=207&rft.pages=40-58&rft.date=2015-06&rft_id=info%3Adoi%2F10.1016%2Fj.jconrel.2015.03.036&rft_id=info%3Apmid%2F25841699&rft.aulast=Milane&rft.aufirst=L&rft.au=Trivedi%2C+M&rft.au=Singh%2C+A&rft.au=Talekar%2C+M&rft.au=Amiji%2C+M&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> <li id="cite_note-228"><a href="#cite_ref-228">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMartinGargZimorski2015" class="citation journal cs1">Martin WF, Garg S, Zimorski V (September 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4571569">"Endosymbiotic theories for eukaryote origin"</a>. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 370 (1678): 20140330. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.2014.0330">10.1098/rstb.2014.0330</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4571569">4571569</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/26323761">26323761</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+of+London.+Series+B%2C+Biological+Sciences&rft.atitle=Endosymbiotic+theories+for+eukaryote+origin&rft.volume=370&rft.issue=1678&rft.pages=20140330&rft.date=2015-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4571569%23id-name%3DPMC&rft_id=info%3Apmid%2F26323761&rft_id=info%3Adoi%2F10.1098%2Frstb.2014.0330&rft.aulast=Martin&rft.aufirst=WF&rft.au=Garg%2C+S&rft.au=Zimorski%2C+V&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4571569&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"> </li> </ol></div> General <ul><li><img alt="Public Domain" src="//upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/12px-PD-icon.svg.png" decoding="async" width="12" height="12" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/18px-PD-icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/62/PD-icon.svg/24px-PD-icon.svg.png 2x" data-file-width="196" data-file-height="196" /> This article incorporates <a href="/wiki/Copyright_status_of_works_by_the_federal_government_of_the_United_States" title="Copyright status of works by the federal government of the United States">public domain material</a> from <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20091208004309/https://www.ncbi.nlm.nih.gov/About/primer/index.html">Science Primer</a>. <a href="/wiki/NCBI" class="mw-redirect" title="NCBI">NCBI</a>. 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The Vital Question: Energy, Evolution, and the Origins of Complex Life. WW Norton & Company. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0393352979" title="Special:BookSources/978-0393352979"><bdi>978-0393352979</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Vital+Question%3A+Energy%2C+Evolution%2C+and+the+Origins+of+Complex+Life&rft.pub=WW+Norton+%26+Company&rft.date=2016&rft.isbn=978-0393352979&rft.aulast=Lane&rft.aufirst=N&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrion" class="Z3988"></li> <li><a rel="nofollow" class="external text" href="https://xvivo.com/examples/powering-the-cell-mitochondria-2/">Powering the Cell Mitochondria</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20220817083144/https://xvivo.com/examples/powering-the-cell-mitochondria-2/">Archived</a> August 17, 2022, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> – XVIVO Scientific Animation</li> <li><a rel="nofollow" class="external text" href="http://www.Mitodb.com">Mitodb.com</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20130703105544/http://www.mitodb.com/">Archived</a> July 3, 2013, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> – The mitochondrial disease database.</li> <li><a rel="nofollow" class="external text" href="http://www.uni-mainz.de/FB/Medizin/Anatomie/workshop/EM/EMMitoE.html">Mitochondria Atlas</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20120629101035/http://www.uni-mainz.de/FB/Medizin/Anatomie/workshop/EM/EMMitoE.html">Archived</a> June 29, 2012, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> at <a href="/wiki/University_of_Mainz" title="University of Mainz">University of Mainz</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080821114550/http://www.mitochondrial.net/">Mitochondria Research Portal</a> at mitochondrial.net</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100125062948/http://www.cytochemistry.net/Cell-biology/mitoch1.htm">Mitochondria: Architecture dictates function</a> at cytochemistry.net</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090418174136/http://bama.ua.edu/~hsmithso/class/bsc_495/mito-plastids/mito_web.html">Mitochondria links</a> at <a href="/wiki/University_of_Alabama" title="University of Alabama">University of Alabama</a></li> <li><a rel="nofollow" class="external text" href="http://www.mitophysiology.org/">MIP</a> <a rel="nofollow" class="external text" href="http://arquivo.pt/wayback/20160523200848/http://www.mitophysiology.org/">Archived</a> May 23, 2016, at the Portuguese Web Archive Mitochondrial Physiology Society</li> <li><a rel="nofollow" class="external text" href="http://opm.phar.umich.edu/localization.php?localization=Mitochondrial%20inner%20membrane">3D structures of proteins from inner mitochondrial membrane</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110913122941/http://opm.phar.umich.edu/localization.php?localization=Mitochondrial%20inner%20membrane">Archived</a> September 13, 2011, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> at <a href="/wiki/University_of_Michigan" title="University of Michigan">University of Michigan</a></li> <li><a rel="nofollow" class="external text" href="http://opm.phar.umich.edu/localization.php?localization=Mitochondrial%20outer%20membrane">3D structures of proteins associated with outer mitochondrial membrane</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110913122952/http://opm.phar.umich.edu/localization.php?localization=Mitochondrial%20outer%20membrane">Archived</a> September 13, 2011, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> at <a href="/wiki/University_of_Michigan" title="University of Michigan">University of Michigan</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20171201205358/http://www.mitoproteins.org/">Mitochondrial Protein Partnership</a> at <a href="/wiki/University_of_Wisconsin" class="mw-redirect" title="University of Wisconsin">University of Wisconsin</a></li> <li><a rel="nofollow" class="external text" href="http://mitominer.mrc-mbu.cam.ac.uk">MitoMiner – A mitochondrial proteomics database</a>[<a href="/wiki/Wikipedia:Link_rot" title="Wikipedia:Link rot">permanent dead link</a>‍] at <a href="/wiki/MRC_Mitochondrial_Biology_Unit" title="MRC Mitochondrial Biology Unit">MRC Mitochondrial Biology Unit</a></li> <li><a rel="nofollow" class="external text" href="http://ccdb.ucsd.edu/sand/main?stype=lite&keyword=mitochondrion&Submit=Go&event=display&start=1">Mitochondrion – Cell Centered 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navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fatty_acid_degradation" title="Fatty acid degradation">fatty acid degradation</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/Carnitine_palmitoyltransferase_I" title="Carnitine palmitoyltransferase I">Carnitine palmitoyltransferase I</a></li> <li><a href="/wiki/Long-chain-fatty-acid%E2%80%94CoA_ligase" title="Long-chain-fatty-acid—CoA ligase">Long-chain-fatty-acid—CoA ligase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> metabolism</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/Kynureninase" title="Kynureninase">Kynureninase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Monoamine_neurotransmitter" title="Monoamine neurotransmitter">monoamine neurotransmitter</a> metabolism</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/Monoamine_oxidase" title="Monoamine oxidase">Monoamine oxidase</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/Mitochondrial_intermembrane_space" class="mw-redirect" title="Mitochondrial intermembrane space">Intermembrane space</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/Adenylate_kinase" title="Adenylate kinase">Adenylate kinase</a></li> <li><a href="/wiki/Creatine_kinase" title="Creatine kinase">Creatine kinase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Inner_mitochondrial_membrane" title="Inner mitochondrial membrane">Inner membrane</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%"><a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</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/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Coenzyme Q – cytochrome c reductase</a></li> <li><a href="/wiki/Cytochrome_c" title="Cytochrome c">Cytochrome c</a></li> <li><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a></li> <li><a href="/wiki/Succinate_dehydrogenase" title="Succinate dehydrogenase">Succinate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pyrimidine_metabolism" title="Pyrimidine metabolism">pyrimidine metabolism</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/Dihydroorotate_dehydrogenase" title="Dihydroorotate dehydrogenase">Dihydroorotate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mitochondrial_shuttle" title="Mitochondrial shuttle">mitochondrial shuttle</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/Malate-aspartate_shuttle" class="mw-redirect" title="Malate-aspartate shuttle">Malate-aspartate shuttle</a></li> <li><a href="/wiki/Glycerol_phosphate_shuttle" title="Glycerol phosphate shuttle">Glycerol phosphate shuttle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Steroidogenesis" class="mw-redirect" title="Steroidogenesis">steroidogenesis</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/Cholesterol_side-chain_cleavage_enzyme" title="Cholesterol side-chain cleavage enzyme">Cholesterol side-chain cleavage enzyme</a></li> <li><a href="/wiki/Steroid_11-beta-hydroxylase" class="mw-redirect" title="Steroid 11-beta-hydroxylase">Steroid 11-beta-hydroxylase</a></li> <li><a href="/wiki/Aldosterone_synthase" title="Aldosterone synthase">Aldosterone synthase</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" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Glutamate_aspartate_transporter" class="mw-redirect" title="Glutamate aspartate transporter">Glutamate aspartate transporter</a></li> <li><a href="/wiki/Glycerol-3-phosphate_dehydrogenase" title="Glycerol-3-phosphate dehydrogenase">Glycerol-3-phosphate dehydrogenase</a></li> <li><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a></li> <li><a href="/wiki/Carnitine_palmitoyltransferase_II" title="Carnitine palmitoyltransferase II">Carnitine palmitoyltransferase II</a></li> <li><a href="/wiki/Uncoupling_protein" title="Uncoupling protein">Uncoupling 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/Mitochondrial_matrix" title="Mitochondrial matrix">Matrix</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%"><a href="/wiki/Citric_acid_cycle" title="Citric acid cycle">citric acid cycle</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/Citrate_synthase" title="Citrate synthase">Citrate synthase</a></li> <li><a href="/wiki/Aconitase" title="Aconitase">Aconitase</a></li> <li><a href="/wiki/Isocitrate_dehydrogenase" title="Isocitrate dehydrogenase">Isocitrate dehydrogenase</a></li> <li><a href="/wiki/Oxoglutarate_dehydrogenase_complex" title="Oxoglutarate dehydrogenase complex">Oxoglutarate dehydrogenase complex</a></li> <li><a href="/wiki/Succinyl_coenzyme_A_synthetase" title="Succinyl coenzyme A synthetase">Succinyl coenzyme A synthetase</a></li> <li><a href="/wiki/Fumarase" title="Fumarase">Fumarase</a></li> <li><a href="/wiki/Malate_dehydrogenase" title="Malate dehydrogenase">Malate dehydrogenase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Anaplerotic_reactions" title="Anaplerotic reactions">anaplerotic reactions</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/Aspartate_transaminase" title="Aspartate transaminase">Aspartate transaminase</a></li> <li><a href="/wiki/Glutamate_dehydrogenase" title="Glutamate dehydrogenase">Glutamate dehydrogenase</a></li> <li><a href="/wiki/Pyruvate_dehydrogenase_complex" title="Pyruvate dehydrogenase complex">Pyruvate dehydrogenase complex</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Urea_cycle" title="Urea cycle">urea cycle</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/Carbamoyl_phosphate_synthetase_I" title="Carbamoyl phosphate synthetase I">Carbamoyl phosphate synthetase I</a></li> <li><a href="/wiki/Ornithine_transcarbamylase" title="Ornithine transcarbamylase">Ornithine transcarbamylase</a></li> <li><a href="/wiki/N-Acetylglutamate_synthase" title="N-Acetylglutamate synthase">N-Acetylglutamate synthase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">alcohol metabolism</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/ALDH2" title="ALDH2">ALDH2</a></li></ul> </div></td></tr><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/PMPCB" title="PMPCB">PMPCB</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/to be sorted</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><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Frataxin" title="Frataxin">Frataxin</a></li> <li><a href="/wiki/Mitochondrial_membrane_transport_protein" title="Mitochondrial membrane transport protein">Mitochondrial membrane transport protein</a> <ul><li><a href="/wiki/Mitochondrial_permeability_transition_pore" title="Mitochondrial permeability transition pore">Mitochondrial permeability transition pore</a></li> <li><a href="/wiki/Mitochondrial_carrier" title="Mitochondrial carrier">Mitochondrial carrier</a></li></ul></li> <li><a href="/wiki/Translocator_protein" title="Translocator protein">Translocator 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/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial DNA</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%"><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">Complex I</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/MT-ND1" title="MT-ND1">MT-ND1</a></li> <li><a href="/wiki/MT-ND2" title="MT-ND2">MT-ND2</a></li> <li><a href="/wiki/MT-ND3" title="MT-ND3">MT-ND3</a></li> <li><a href="/wiki/MT-ND4" title="MT-ND4">MT-ND4</a></li> <li><a href="/wiki/MT-ND4L" title="MT-ND4L">MT-ND4L</a></li> <li><a href="/wiki/MT-ND5" title="MT-ND5">MT-ND5</a></li> <li><a href="/wiki/MT-ND6" title="MT-ND6">MT-ND6</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Coenzyme_Q_%E2%80%93_cytochrome_c_reductase" title="Coenzyme Q – cytochrome c reductase">Complex III</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/MT-CYB" title="MT-CYB">MT-CYB</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Complex IV</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/Cytochrome_c_oxidase_subunit_I" title="Cytochrome c oxidase subunit I">MT-CO1</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_II" class="mw-redirect" title="Cytochrome c oxidase subunit II">MT-CO2</a></li> <li><a href="/wiki/Cytochrome_c_oxidase_subunit_III" title="Cytochrome c oxidase subunit III">MT-CO3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</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/MT-ATP6" title="MT-ATP6">MT-ATP6</a></li> <li><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transfer_RNA" title="Transfer RNA">tRNA</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/MT-TA" title="MT-TA">MT-TA</a></li> <li><a href="/wiki/MT-TC" title="MT-TC">MT-TC</a></li> <li><a href="/wiki/MT-TD" title="MT-TD">MT-TD</a></li> <li><a href="/wiki/MT-TE" title="MT-TE">MT-TE</a></li> <li><a href="/wiki/MT-TF" title="MT-TF">MT-TF</a></li> <li><a href="/wiki/MT-TG" title="MT-TG">MT-TG</a></li> <li><a href="/wiki/MT-TH" title="MT-TH">MT-TH</a></li> <li><a href="/wiki/MT-TI" title="MT-TI">MT-TI</a></li> <li><a href="/wiki/MT-TK" title="MT-TK">MT-TK</a></li> <li><a href="/wiki/MT-TL1" title="MT-TL1">MT-TL1</a></li> <li><a href="/wiki/MT-TL2" title="MT-TL2">MT-TL2</a></li> <li><a href="/wiki/MT-TM" title="MT-TM">MT-TM</a></li> <li><a href="/wiki/MT-TN" title="MT-TN">MT-TN</a></li> <li><a href="/wiki/MT-TP" title="MT-TP">MT-TP</a></li> <li><a href="/wiki/MT-TQ" title="MT-TQ">MT-TQ</a></li> <li><a href="/wiki/MT-TR" title="MT-TR">MT-TR</a></li> <li><a href="/wiki/MT-TS1" title="MT-TS1">MT-TS1</a></li> <li><a href="/wiki/MT-TS2" title="MT-TS2">MT-TS2</a></li> <li><a href="/wiki/MT-TT" title="MT-TT">MT-TT</a></li> <li><a href="/wiki/MT-TV_(mitochondrial)" title="MT-TV (mitochondrial)">MT-TV</a></li> <li><a href="/wiki/MT-TW" title="MT-TW">MT-TW</a></li> <li><a href="/wiki/MT-TY" title="MT-TY">MT-TY</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div>see also <a href="/wiki/Template:Mitochondrial_diseases" title="Template:Mitochondrial diseases">mitochondrial diseases</a></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="Structures_of_the_cell_/_organelles" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cellular_structures" title="Template:Cellular structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cellular_structures" title="Template talk:Cellular structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cellular_structures" title="Special:EditPage/Template:Cellular structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Structures_of_the_cell_/_organelles" style="font-size:114%;margin:0 4em">Structures of the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> / <a href="/wiki/Organelle" title="Organelle">organelles</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endomembrane_system" title="Endomembrane system">Endomembrane system</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_membrane" title="Cell membrane">Cell membrane</a></li> <li><a href="/wiki/Cell_nucleus" title="Cell nucleus">Nucleus</a></li> <li><a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">Endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a></li> <li><a href="/wiki/Parenthesome" title="Parenthesome">Parenthesome</a></li> <li><a href="/wiki/Autophagosome" title="Autophagosome">Autophagosome</a></li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a> <ul><li><a href="/wiki/Exosome_(vesicle)" title="Exosome (vesicle)">Exosome</a></li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Endosome" title="Endosome">Endosome</a></li> <li><a href="/wiki/Phagosome" title="Phagosome">Phagosome</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Acrosome" title="Acrosome">Acrosome</a></li></ul></li> <li><a href="/wiki/Granule_(cell_biology)" title="Granule (cell biology)">Cytoplasmic granule</a> <ul><li><a href="/wiki/Melanosome" title="Melanosome">Melanosome</a></li> <li><a href="/wiki/Microbody" title="Microbody">Microbody</a></li> <li><a href="/wiki/Glyoxysome" title="Glyoxysome">Glyoxysome</a></li> <li><a href="/wiki/Peroxisome" title="Peroxisome">Peroxisome</a></li> <li><a href="/wiki/Weibel%E2%80%93Palade_body" title="Weibel–Palade body">Weibel–Palade body</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Microfilament" title="Microfilament">Microfilament</a></li> <li><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate filament</a></li> <li><a href="/wiki/Microtubule" title="Microtubule">Microtubule</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a></li> <li><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organizing center</a> <ul><li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Centriole" title="Centriole">Centriole</a></li> <li><a href="/wiki/Basal_body" title="Basal body">Basal body</a></li> <li><a href="/wiki/Spindle_pole_body" title="Spindle pole body">Spindle pole body</a></li></ul></li> <li><a href="/wiki/Myofibril" title="Myofibril">Myofibril</a></li> <li><a href="/wiki/Undulipodium" title="Undulipodium">Undulipodium</a> <ul><li><a href="/wiki/Cilium" title="Cilium">Cilium</a></li> <li><a href="/wiki/Flagellum#Eukaryotic" title="Flagellum">Flagellum</a></li> <li><a href="/wiki/Axoneme" title="Axoneme">Axoneme</a></li> <li><a href="/wiki/Radial_spoke" title="Radial spoke">Radial spoke</a></li></ul></li> <li><a href="/wiki/Pseudopodium" class="mw-redirect" title="Pseudopodium">Pseudopodium</a> <ul><li><a href="/wiki/Lamellipodium" title="Lamellipodium">Lamellipodium</a></li> <li><a href="/wiki/Filopodium" class="mw-redirect" title="Filopodium">Filopodium</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbionts</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Amyloplast" title="Amyloplast">Amyloplast</a></li> <li><a href="/wiki/Elaioplast" title="Elaioplast">Elaioplast</a></li> <li><a href="/wiki/Proteinoplast" title="Proteinoplast">Proteinoplast</a></li> <li><a href="/wiki/Tannosome" title="Tannosome">Tannosome</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other internal</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Spliceosome" title="Spliceosome">Spliceosome</a></li> <li><a href="/wiki/Vault_(organelle)" title="Vault (organelle)">Vault</a></li></ul></li> <li><a href="/wiki/Cytoplasm" title="Cytoplasm">Cytoplasm</a> <ul><li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a></li> <li><a href="/wiki/Cytoplasmic_inclusion" class="mw-redirect" title="Cytoplasmic inclusion">Inclusions</a></li></ul></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</a></li> <li><a href="/wiki/Magnetosome" title="Magnetosome">Magnetosome</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">External</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Extracellular_matrix" title="Extracellular matrix">Extracellular matrix</a></li></ul> </div></td></tr></tbody></table></div> <div 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="Life,_non-cellular_life,_and_comparable_structures" 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:Life,_non-cellular_life,_and_comparable_structures" title="Template:Life, non-cellular life, and comparable structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life,_non-cellular_life,_and_comparable_structures" title="Template talk:Life, non-cellular life, and comparable structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life,_non-cellular_life,_and_comparable_structures" title="Special:EditPage/Template:Life, non-cellular life, and comparable structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Life,_non-cellular_life,_and_comparable_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Life" title="Life">Life</a>, <a href="/wiki/Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</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/Prokaryote" title="Prokaryote">Prokaryota</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/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> <ul><li><a class="mw-selflink selflink">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li></ul></li> <li><a href="/wiki/LUCA" class="mw-redirect" title="LUCA">LUCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</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/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li>'<a href="/wiki/Protist" title="Protist">Protista</a>'</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</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/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</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/Virus" title="Virus">Virus</a> <hr /><a href="/wiki/Viroid" title="Viroid">Viroid</a> <hr /><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</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/Adnaviria" title="Adnaviria">Adnaviria</a></li> <li><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></li> <li><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></li> <li><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></li> <li><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></li> <li><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</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/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></li> <li><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></li> <li><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li> <li><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></li> <li><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></li> <li><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></li> <li><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></li> <li><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></li> <li><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></li> <li><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></li> <li><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></li> <li><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></li> <li><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></li> <li><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></li> <li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></li> <li><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></li> <li><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</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/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></li> <li><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></li></ul> </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%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nanobe" title="Nanobe">Nanobe</a> (?)</li> <li><a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable structures</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/Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li> <li>"<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a>"</li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Prion" title="Prion">Prion</a> <ul><li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a> <a href="/wiki/Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li> <li>Retroelements not elsewhere classified <ul><li><a href="/wiki/Retron" title="Retron">Retron</a></li> <li><a href="/wiki/Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li> <li><a href="/wiki/Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li> <li><a href="/wiki/Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Spiegelman%27s_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li> <li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retroposon" title="Retroposon">Retroposon</a></li></ul></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li> <li><a href="/wiki/Xenobot" title="Xenobot">Xenobot</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="Self-replicating_organic_structures" 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="text-align: center;"><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:Self-replicating_organic_structures" title="Template:Self-replicating organic structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Self-replicating_organic_structures" title="Template talk:Self-replicating organic structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Self-replicating_organic_structures" title="Special:EditPage/Template:Self-replicating organic structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Self-replicating_organic_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</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/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</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/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral agents</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:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="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%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</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>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</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/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a> self-replication</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:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</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 class="mw-selflink selflink">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</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/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</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/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" 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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