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Replication </div> </a> <ul id="toc-Replication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_on_the_human_mtDNA_and_their_transcription" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genes_on_the_human_mtDNA_and_their_transcription"> <div class="vector-toc-text"> 4 Genes on the human mtDNA and their transcription </div> </a> <button aria-controls="toc-Genes_on_the_human_mtDNA_and_their_transcription-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Genes on the human mtDNA and their transcription subsection </button> <ul id="toc-Genes_on_the_human_mtDNA_and_their_transcription-sublist" class="vector-toc-list"> <li id="toc-Regulation_of_transcription" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Regulation_of_transcription"> <div class="vector-toc-text"> 4.1 Regulation of transcription </div> </a> <ul id="toc-Regulation_of_transcription-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mitochondrial_inheritance" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mitochondrial_inheritance"> <div class="vector-toc-text"> 5 Mitochondrial inheritance </div> </a> <button aria-controls="toc-Mitochondrial_inheritance-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Mitochondrial inheritance subsection </button> <ul id="toc-Mitochondrial_inheritance-sublist" class="vector-toc-list"> <li id="toc-Female_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Female_inheritance"> <div class="vector-toc-text"> 5.1 Female inheritance </div> </a> <ul id="toc-Female_inheritance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-The_mitochondrial_bottleneck" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_mitochondrial_bottleneck"> <div class="vector-toc-text"> 5.2 The mitochondrial bottleneck </div> </a> <ul id="toc-The_mitochondrial_bottleneck-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Male_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Male_inheritance"> <div class="vector-toc-text"> 5.3 Male inheritance </div> </a> <ul id="toc-Male_inheritance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mitochondrial_donation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitochondrial_donation"> <div class="vector-toc-text"> 5.4 Mitochondrial donation </div> </a> <ul id="toc-Mitochondrial_donation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mutations_and_disease" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mutations_and_disease"> <div class="vector-toc-text"> 6 Mutations and disease </div> </a> <button aria-controls="toc-Mutations_and_disease-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Mutations and disease subsection </button> <ul id="toc-Mutations_and_disease-sublist" class="vector-toc-list"> <li id="toc-Susceptibility" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Susceptibility"> <div class="vector-toc-text"> 6.1 Susceptibility </div> </a> <ul id="toc-Susceptibility-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_illness" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_illness"> <div class="vector-toc-text"> 6.2 Genetic illness </div> </a> <ul id="toc-Genetic_illness-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Use_in_disease_diagnosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Use_in_disease_diagnosis"> <div class="vector-toc-text"> 6.3 Use in disease diagnosis </div> </a> <ul id="toc-Use_in_disease_diagnosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Relationship_with_ageing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Relationship_with_ageing"> <div class="vector-toc-text"> 6.4 Relationship with ageing </div> </a> <ul id="toc-Relationship_with_ageing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neurodegenerative_diseases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neurodegenerative_diseases"> <div class="vector-toc-text"> 6.5 Neurodegenerative diseases </div> </a> <ul id="toc-Neurodegenerative_diseases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Correlation_of_the_mtDNA_base_composition_with_animal_life_spans" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Correlation_of_the_mtDNA_base_composition_with_animal_life_spans"> <div class="vector-toc-text"> 6.6 Correlation of the mtDNA base composition with animal life spans </div> </a> <ul id="toc-Correlation_of_the_mtDNA_base_composition_with_animal_life_spans-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-mtDNA_mutational_spectrum_is_sensitive_to_species-specific_life-history_traits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#mtDNA_mutational_spectrum_is_sensitive_to_species-specific_life-history_traits"> <div class="vector-toc-text"> 6.7 mtDNA mutational spectrum is sensitive to species-specific life-history traits </div> </a> <ul id="toc-mtDNA_mutational_spectrum_is_sensitive_to_species-specific_life-history_traits-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Relationship_with_non-B_(non-canonical)_DNA_structures" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Relationship_with_non-B_(non-canonical)_DNA_structures"> <div class="vector-toc-text"> 6.8 Relationship with non-B (non-canonical) DNA structures </div> </a> <ul id="toc-Relationship_with_non-B_(non-canonical)_DNA_structures-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Use_in_forensics" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Use_in_forensics"> <div class="vector-toc-text"> 7 Use in forensics </div> </a> <ul id="toc-Use_in_forensics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Use_in_evolutionary_biology_and_systematic_biology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Use_in_evolutionary_biology_and_systematic_biology"> <div class="vector-toc-text"> 8 Use in evolutionary biology and systematic biology </div> </a> <ul id="toc-Use_in_evolutionary_biology_and_systematic_biology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-mtDNA_in_nuclear_DNA" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#mtDNA_in_nuclear_DNA"> <div class="vector-toc-text"> 9 mtDNA in nuclear DNA </div> </a> <ul id="toc-mtDNA_in_nuclear_DNA-sublist" class="vector-toc-list"> </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"> 10 History </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mitochondrial_sequence_databases" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mitochondrial_sequence_databases"> <div class="vector-toc-text"> 11 Mitochondrial sequence databases </div> </a> <ul id="toc-Mitochondrial_sequence_databases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-MtDNA-phenotype_association_databases" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#MtDNA-phenotype_association_databases"> <div class="vector-toc-text"> 12 MtDNA-phenotype association databases </div> </a> <button aria-controls="toc-MtDNA-phenotype_association_databases-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle MtDNA-phenotype association databases subsection </button> <ul id="toc-MtDNA-phenotype_association_databases-sublist" class="vector-toc-list"> <li id="toc-Mitochondrial_mutation_databases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mitochondrial_mutation_databases"> <div class="vector-toc-text"> 12.1 Mitochondrial mutation databases </div> </a> <ul id="toc-Mitochondrial_mutation_databases-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 13 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"> 14 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"> 15 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">Mitochondrial DNA</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 54 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-54" aria-hidden="true" > 54 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/Mitochondriale_DNS" title="Mitochondriale DNS – Afrikaans" lang="af" hreflang="af" data-title="Mitochondriale DNS" 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/%D8%AF%D9%86%D8%A7_%D9%85%D8%AA%D9%82%D8%AF%D8%B1%D8%A9" title="دنا متقدرة – Arabic" lang="ar" hreflang="ar" data-title="دنا متقدرة" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</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_DNA" title="Soàⁿ-lia̍p-thé DNA – Minnan" lang="nan" hreflang="nan" data-title="Soàⁿ-lia̍p-thé DNA" 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%B0%D0%BD%D0%B4%D1%80%D1%8B%D1%8F%D0%BB%D1%8C%D0%BD%D0%B0%D1%8F_%D0%94%D0%9D%D0%9A" title="Мітахандрыяльная ДНК – Belarusian" lang="be" hreflang="be" data-title="Мітахандрыяльная ДНК" 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%D0%B0%D0%BB%D0%BD%D0%B0_%D0%94%D0%9D%D0%9A" title="Митохондриална ДНК – Bulgarian" lang="bg" hreflang="bg" data-title="Митохондриална ДНК" 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/Mitohondrijska_DNK" title="Mitohondrijska DNK – Bosnian" lang="bs" hreflang="bs" data-title="Mitohondrijska DNK" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/ADN_mitocondrial" title="ADN mitocondrial – Catalan" lang="ca" hreflang="ca" data-title="ADN mitocondrial" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Mitochondri%C3%A1ln%C3%AD_DNA" title="Mitochondriální DNA – Czech" lang="cs" hreflang="cs" data-title="Mitochondriální DNA" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Mitokondrielt_DNA" title="Mitokondrielt DNA – Danish" lang="da" hreflang="da" data-title="Mitokondrielt DNA" 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/Mitochondriale_DNA" title="Mitochondriale DNA – German" lang="de" hreflang="de" data-title="Mitochondriale DNA" 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/Mitokondriaalne_DNA" title="Mitokondriaalne DNA – Estonian" lang="et" hreflang="et" data-title="Mitokondriaalne DNA" 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%CE%BF%CE%BD%CE%B4%CF%81%CE%B9%CE%B1%CE%BA%CF%8C_DNA" title="Μιτοχονδριακό DNA – Greek" lang="el" hreflang="el" data-title="Μιτοχονδριακό DNA" 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/Genoma_mitocondrial" title="Genoma mitocondrial – Spanish" lang="es" hreflang="es" data-title="Genoma mitocondrial" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/DNA_mitokondrial" title="DNA mitokondrial – Basque" lang="eu" hreflang="eu" data-title="DNA mitokondrial" 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/%D8%AF%DB%8C%E2%80%8C%D8%A7%D9%86%E2%80%8C%D8%A7%DB%8C_%D9%85%DB%8C%D8%AA%D9%88%DA%A9%D9%86%D8%AF%D8%B1%DB%8C%D8%A7%DB%8C%DB%8C" title="دی‌ان‌ای میتوکندریایی – Persian" lang="fa" hreflang="fa" data-title="دی‌ان‌ای میتوکندریایی" 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/G%C3%A9nome_mitochondrial" title="Génome mitochondrial – French" lang="fr" hreflang="fr" data-title="Génome mitochondrial" 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/Mitogondriaal_DNA" title="Mitogondriaal DNA – Western Frisian" lang="fy" hreflang="fy" data-title="Mitogondriaal DNA" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target">Frysk</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/ADN_mitocondrial" title="ADN mitocondrial – Galician" lang="gl" hreflang="gl" data-title="ADN mitocondrial" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</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_DNA" title="미토콘드리아 DNA – Korean" lang="ko" hreflang="ko" data-title="미토콘드리아 DNA" 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%A1%D5%AC_%D4%B4%D5%86%D4%B9" title="Միտոքոնդրիալ ԴՆԹ – Armenian" lang="hy" hreflang="hy" data-title="Միտոքոնդրիալ ԴՆԹ" 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%AE%E0%A4%BE%E0%A4%87%E0%A4%9F%E0%A5%8B%E0%A4%95%E0%A4%BE%E0%A4%A3%E0%A5%8D%E0%A4%A1%E0%A5%8D%E0%A4%B0%E0%A4%BF%E0%A4%AF%E0%A4%BE%E0%A4%88_%E0%A4%A1%E0%A5%80%E0%A4%8F%E0%A4%A8%E0%A4%8F" title="माइटोकाण्ड्रियाई डीएनए – Hindi" lang="hi" hreflang="hi" data-title="माइटोकाण्ड्रियाई डीएनए" 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/Mitohondrijski_DNK" title="Mitohondrijski DNK – Croatian" lang="hr" hreflang="hr" data-title="Mitohondrijski DNK" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/DNA_mitokondria" title="DNA mitokondria – Indonesian" lang="id" hreflang="id" data-title="DNA mitokondria" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/DNA_mitocondriale" title="DNA mitocondriale – Italian" lang="it" hreflang="it" data-title="DNA mitocondriale" 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%92%D7%A0%D7%95%D7%9D_%D7%9E%D7%99%D7%98%D7%95%D7%9B%D7%95%D7%A0%D7%93%D7%A8%D7%99" title="גנום מיטוכונדרי – Hebrew" lang="he" hreflang="he" data-title="גנום מיטוכונדרי" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</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%D0%BB%D1%8B%D2%9B_%D0%94%D0%9D%D2%9A" title="Митохондриялық ДНҚ – Kazakh" lang="kk" hreflang="kk" data-title="Митохондриялық ДНҚ" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/DNA_ya_mitokondria" title="DNA ya mitokondria – Swahili" lang="sw" hreflang="sw" data-title="DNA ya mitokondria" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target">Kiswahili</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/ADN_mitochondriale" title="ADN mitochondriale – Latin" lang="la" hreflang="la" data-title="ADN mitochondriale" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</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%81%D0%BA%D0%B0_%D0%94%D0%9D%D0%9A" title="Митохондриска ДНК – Macedonian" lang="mk" hreflang="mk" data-title="Митохондриска ДНК" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/DNA_mitokondrion" title="DNA mitokondrion – Malay" lang="ms" hreflang="ms" data-title="DNA mitokondrion" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9C%D0%B8%D1%82%D0%BE%D1%85%D0%BE%D0%BD%D0%B4%D1%80%D0%B8%D0%B9%D0%BD_%D0%94%D0%9D%D0%A5" title="Митохондрийн ДНХ – Mongolian" lang="mn" hreflang="mn" data-title="Митохондрийн ДНХ" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target">Монгол</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Mitochondriaal_DNA" title="Mitochondriaal DNA – Dutch" lang="nl" hreflang="nl" data-title="Mitochondriaal DNA" 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%A2DNA" title="ミトコンドリアDNA – Japanese" lang="ja" hreflang="ja" data-title="ミトコンドリアDNA" 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 badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://frr.wikipedia.org/wiki/Mitochondriol_DNA" title="Mitochondriol DNA – Northern Frisian" lang="frr" hreflang="frr" data-title="Mitochondriol DNA" 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/Mitokondrielt_DNA" title="Mitokondrielt DNA – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Mitokondrielt DNA" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%85%D8%A7%DB%8C%D8%AA%D9%88%DA%A9%D8%A7%D9%86%D8%AF%D8%B1%D9%8A_%DA%89%D9%8A.%D8%A7%DB%90%D9%86.%D8%A7%DB%90" title="مایتوکاندري ډي.اېن.اې – Pashto" lang="ps" hreflang="ps" data-title="مایتوکاندري ډي.اېن.اې" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Mitochondrialny_DNA" title="Mitochondrialny DNA – Polish" lang="pl" hreflang="pl" data-title="Mitochondrialny DNA" 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/ADN_mitocondrial" title="ADN mitocondrial – Portuguese" lang="pt" hreflang="pt" data-title="ADN mitocondrial" 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/ADN_mitocondrial" title="ADN mitocondrial – Romanian" lang="ro" hreflang="ro" data-title="ADN mitocondrial" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</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%D0%B0%D0%BB%D1%8C%D0%BD%D0%B0%D1%8F_%D0%94%D0%9D%D0%9A" title="Митохондриальная ДНК – Russian" lang="ru" hreflang="ru" data-title="Митохондриальная ДНК" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-stq mw-list-item"><a href="https://stq.wikipedia.org/wiki/Mitochondrioale_DNA" title="Mitochondrioale DNA – Saterland Frisian" lang="stq" hreflang="stq" data-title="Mitochondrioale DNA" data-language-autonym="Seeltersk" data-language-local-name="Saterland Frisian" class="interlanguage-link-target">Seeltersk</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Mitochondrial_DNA" title="Mitochondrial DNA – Simple English" lang="en-simple" hreflang="en-simple" data-title="Mitochondrial DNA" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sd mw-list-item"><a href="https://sd.wikipedia.org/wiki/%D9%85%D8%A7%D8%A6%D9%8A%D9%BD%D9%88%DA%AA%D9%88%D9%86%DA%8A%D8%B1%D9%8A%D9%84_%DA%8A%D9%8A_%D8%A7%D9%8A%D9%86_%D8%A7%D9%8A" title="مائيٽوڪونڊريل ڊي اين اي – Sindhi" lang="sd" hreflang="sd" data-title="مائيٽوڪونڊريل ڊي اين اي" data-language-autonym="سنڌي" data-language-local-name="Sindhi" class="interlanguage-link-target">سنڌي</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Mitochondri%C3%A1lna_DNA" title="Mitochondriálna DNA – Slovak" lang="sk" hreflang="sk" data-title="Mitochondriálna DNA" 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/Mitohondrijska_DNK" title="Mitohondrijska DNK – Slovenian" lang="sl" hreflang="sl" data-title="Mitohondrijska DNK" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</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%D1%81%D0%BA%D0%B8_%D0%B3%D0%B5%D0%BD%D0%BE%D0%BC" title="Митохондријски геном – Serbian" lang="sr" hreflang="sr" data-title="Митохондријски геном" 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/Mitohondrijski_genom" title="Mitohondrijski genom – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Mitohondrijski genom" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Mitokondriellt_DNA" title="Mitokondriellt DNA – Swedish" lang="sv" hreflang="sv" data-title="Mitokondriellt DNA" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</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%E0%AE%95%E0%AE%B3%E0%AE%BF%E0%AE%A9%E0%AF%8D_%E0%AE%9F%E0%AE%BF_%E0%AE%86%E0%AE%95%E0%AF%8D%E0%AE%9A%E0%AE%BF-%E0%AE%B0%E0%AF%88%E0%AE%AA%E0%AF%8B%E0%AE%A8%E0%AE%BF%E0%AE%AF%E0%AF%82%E0%AE%95%E0%AF%8D%E0%AE%B2%E0%AE%BF%E0%AE%AF%E0%AE%BF%E0%AE%95%E0%AF%8D_%E0%AE%95%E0%AE%BE%E0%AE%9F%E0%AE%BF" title="இழைமணிகளின் டி ஆக்சி-ரைபோநியூக்லியிக் காடி – Tamil" lang="ta" hreflang="ta" data-title="இழைமணிகளின் டி ஆக்சி-ரைபோநியூக்லியிக் காடி" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%94%E0%B8%B5%E0%B9%80%E0%B8%AD%E0%B9%87%E0%B8%99%E0%B9%80%E0%B8%AD%E0%B8%82%E0%B8%AD%E0%B8%87%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="ดีเอ็นเอของไมโทคอนเดรีย – Thai" lang="th" hreflang="th" data-title="ดีเอ็นเอของไมโทคอนเดรีย" 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/Mitokondriyal_DNA" title="Mitokondriyal DNA – Turkish" lang="tr" hreflang="tr" data-title="Mitokondriyal DNA" 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%D0%B0%D0%BB%D1%8C%D0%BD%D0%B0_%D0%94%D0%9D%D0%9A" title="Мітохондріальна ДНК – Ukrainian" lang="uk" hreflang="uk" data-title="Мітохондріальна ДНК" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/DNA_ty_th%E1%BB%83" title="DNA ty thể – Vietnamese" lang="vi" hreflang="vi" data-title="DNA ty thể" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%BA%BF%E7%B2%92%E4%BD%93DNA" title="线粒体DNA – Chinese" lang="zh" hreflang="zh" data-title="线粒体DNA" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li> 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.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">For the journal, see <a href="/wiki/Mitochondrial_DNA_(journal)" title="Mitochondrial DNA (journal)">Mitochondrial DNA (journal)</a>.</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Mitochondrial_DNA_lg.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Mitochondrial_DNA_lg.jpg/300px-Mitochondrial_DNA_lg.jpg" decoding="async" width="300" height="335" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Mitochondrial_DNA_lg.jpg/450px-Mitochondrial_DNA_lg.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/17/Mitochondrial_DNA_lg.jpg/600px-Mitochondrial_DNA_lg.jpg 2x" data-file-width="800" data-file-height="894" /></a><figcaption>Mitochondrial DNA is the small <a href="/wiki/Circular_chromosome" title="Circular chromosome">circular chromosome</a> found inside mitochondria. These <a href="/wiki/Organelle" title="Organelle">organelles</a>, found in all <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> cells, are the powerhouse of the cell.<a href="#cite_note-Siekevitz-1">[1]</a> The mitochondria, and thus mitochondrial DNA, are passed exclusively from <a href="/wiki/Mother" title="Mother">mother</a> to <a href="/wiki/Offspring" title="Offspring">offspring</a> through the <a href="/wiki/Egg_cell" title="Egg cell">egg cell</a>.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/Mitochondrial_DNA_3D.webm/220px--Mitochondrial_DNA_3D.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="165" data-durationhint="33" data-mwtitle="Mitochondrial_DNA_3D.webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Mitochondrial_DNA_3D.webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/2/25/Mitochondrial_DNA_3D.webm/Mitochondrial_DNA_3D.webm.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="640" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/2/25/Mitochondrial_DNA_3D.webm" type="video/webm; codecs="vp9, vorbis"" data-width="640" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/2/25/Mitochondrial_DNA_3D.webm/Mitochondrial_DNA_3D.webm.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="192" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/2/25/Mitochondrial_DNA_3D.webm/Mitochondrial_DNA_3D.webm.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="320" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/2/25/Mitochondrial_DNA_3D.webm/Mitochondrial_DNA_3D.webm.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="480" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/2/25/Mitochondrial_DNA_3D.webm/Mitochondrial_DNA_3D.webm.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="480" data-height="360" /></video><figcaption>Illustration of the location of mitochondrial DNA in human cells</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg/220px-Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg" decoding="async" width="220" height="284" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg/330px-Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg/440px-Electron_microscopy_reveals_mitochondrial_DNA_in_discrete_foci.jpg 2x" data-file-width="956" data-file-height="1236" /></a><figcaption>Electron microscopy reveals mitochondrial DNA in discrete foci. Bars: 200 nm. (A) Cytoplasmic section after <a href="/wiki/Immunogold_labelling" title="Immunogold labelling">immunogold labelling</a> with anti-DNA; gold particles marking mtDNA are found near the mitochondrial membrane (black dots in upper right). (B) Whole mount view of cytoplasm after extraction with CSK buffer and immunogold labelling with anti-DNA; mtDNA (marked by gold particles) resists extraction. From Iborra et al., 2004.[<a href="/wiki/Wikipedia:Manual_of_Style#Technical_language" title="Wikipedia:Manual of Style">jargon</a>]<a href="#cite_note-pmid15157274-2">[2]</a></figcaption></figure> Mitochondrial DNA (mtDNA and mDNA) is the <a href="/wiki/DNA" title="DNA">DNA</a> located in the <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> <a href="/wiki/Organelle" title="Organelle">organelles</a> in a <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> cell that converts chemical energy from food into <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP). Mitochondrial DNA is a small portion of the DNA contained in a eukaryotic cell; most of the DNA is in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>, and, in plants and algae, the DNA also is found in <a href="/wiki/Plastid" title="Plastid">plastids</a>, such as <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a>.<a href="#cite_note-3">[3]</a> <a href="/wiki/Human_mitochondrial_DNA" class="mw-redirect" title="Human mitochondrial DNA">Human mitochondrial DNA</a> was the first significant part of the <a href="/wiki/Human_genome" title="Human genome">human genome</a> to be sequenced.<a href="#cite_note-pmid7219534-4">[4]</a> This sequencing revealed that human mtDNA has 16,569 <a href="/wiki/Base_pair" title="Base pair">base pairs</a> and encodes 13 <a href="/wiki/Protein" title="Protein">proteins</a>. As in other vertebrates, the human mitochondrial <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> differs slightly from nuclear DNA.<a href="#cite_note-5">[5]</a> Since animal mtDNA evolves faster than <a href="/wiki/Nuclear_gene" title="Nuclear gene">nuclear genetic</a> markers,<a href="#cite_note-6">[6]</a><a href="#cite_note-7">[7]</a><a href="#cite_note-8">[8]</a> it represents a mainstay of <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogenetics</a> and <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>. It also permits tracing the relationships of populations, and so has become important in <a href="/wiki/Anthropology" title="Anthropology">anthropology</a> and <a href="/wiki/Biogeography" title="Biogeography">biogeography</a>. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Origin">Origin</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=1" title="Edit section: Origin">edit</a>]</div> Nuclear and mitochondrial DNA are thought to have separate <a href="/wiki/Evolution" title="Evolution">evolutionary</a> origins, with the mtDNA derived from the circular genomes of <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> engulfed by the ancestors of modern eukaryotic cells. This theory is called the <a href="/wiki/Endosymbiotic_theory" class="mw-redirect" title="Endosymbiotic theory">endosymbiotic theory</a>. In the cells of extant organisms, the vast majority of the proteins in the mitochondria (numbering approximately 1500 different types in <a href="/wiki/Mammal" title="Mammal">mammals</a>) are coded by <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a>, but the genes for some, if not most, of them are thought to be of bacterial origin, having been transferred to the <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> nucleus during <a href="/wiki/Evolution" title="Evolution">evolution</a>.<a href="#cite_note-Johnston,_I._G._and_Williams,_B._P._2016-9">[9]</a> The reasons mitochondria have retained some genes are debated. The existence in some species of mitochondrion-derived organelles lacking a genome<a href="#cite_note-10">[10]</a> suggests that complete gene loss is possible, and transferring mitochondrial genes to the nucleus has several advantages.<a href="#cite_note-11">[11]</a> The difficulty of targeting remotely-produced hydrophobic protein products to the mitochondrion is one hypothesis for why some genes are retained in mtDNA;<a href="#cite_note-12">[12]</a> <a href="/wiki/CoRR_hypothesis" title="CoRR hypothesis">colocalisation for redox regulation</a> is another, citing the desirability of localised control over mitochondrial machinery.<a href="#cite_note-13">[13]</a> Recent analysis of a wide range of mtDNA genomes suggests that both these features may dictate mitochondrial gene retention.<a href="#cite_note-Johnston,_I._G._and_Williams,_B._P._2016-9">[9]</a> <div class="mw-heading mw-heading2"><h2 id="Genome_structure_and_diversity">Genome structure and diversity</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=2" title="Edit section: Genome structure and diversity">edit</a>]</div> Across all organisms, there are six main mitochondrial genome types, classified by structure (i.e. circular versus linear), size, presence of <a href="/wiki/Intron" title="Intron">introns</a> or <a href="/wiki/Plasmid" title="Plasmid">plasmid like structures</a>, and whether the genetic material is a singular molecule or collection of <a href="/wiki/Homogeneity_and_heterogeneity" title="Homogeneity and heterogeneity">homogeneous</a> or <a href="/wiki/Homogeneity_and_heterogeneity" title="Homogeneity and heterogeneity">heterogeneous</a> molecules.<a href="#cite_note-Kolesnikov,_A._A._2012-14">[14]</a> In many unicellular organisms (e.g., the <a href="/wiki/Ciliate" title="Ciliate">ciliate</a> <a href="/wiki/Tetrahymena" title="Tetrahymena">Tetrahymena</a> and the <a href="/wiki/Green_alga" class="mw-redirect" title="Green alga">green alga</a> <a href="/wiki/Chlamydomonas_reinhardtii" title="Chlamydomonas reinhardtii">Chlamydomonas reinhardtii</a>), and in rare cases also in multicellular organisms (e.g. in some species of <a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a>), the mtDNA is linear <a href="/wiki/DNA" title="DNA">DNA</a>. Most of these linear mtDNAs possess <a href="/wiki/Telomerase" title="Telomerase">telomerase</a>-independent <a href="/wiki/Telomere" title="Telomere">telomeres</a> (i.e., the ends of the linear <a href="/wiki/DNA" title="DNA">DNA</a>) with different modes of replication, which have made them interesting objects of research because many of these unicellular organisms with linear mtDNA are known <a href="/wiki/Pathogen" title="Pathogen">pathogens</a>.<a href="#cite_note-15">[15]</a> <div class="mw-heading mw-heading3"><h3 id="Animals">Animals</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=3" title="Edit section: Animals">edit</a>]</div> Most (<a href="/wiki/Bilateria" title="Bilateria">bilaterian</a>) animals have a circular mitochondrial genome. <a href="/wiki/Medusozoa" title="Medusozoa">Medusozoa</a> and <a href="/wiki/Calcareous_sponge" title="Calcareous sponge">calcarea</a> <a href="/wiki/Clade" title="Clade">clades</a> however include species with linear mitochondrial chromosomes.<a href="#cite_note-16">[16]</a> With a few exceptions, animals have 37 genes in their mitochondrial DNA: 13 for <a href="/wiki/Protein" title="Protein">proteins</a>, 22 for <a href="/wiki/Transfer_RNA" title="Transfer RNA">tRNAs</a>, and 2 for <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">rRNAs</a>.<a href="#cite_note-Boore1999-17">[17]</a> Mitochondrial genomes for animals average about 16,000 base pairs in length.<a href="#cite_note-Boore1999-17">[17]</a> The anemone Isarachnanthus nocturnus has the largest mitochondrial genome of any animal at 80,923 bp.<a href="#cite_note-18">[18]</a> The smallest known mitochondrial genome in animals belongs to the comb jelly <a href="/wiki/Vallicula_multiformis" class="mw-redirect" title="Vallicula multiformis">Vallicula multiformis</a>, which consist of 9,961 bp.<a href="#cite_note-19">[19]</a> In February 2020, a jellyfish-related parasite – <a href="/wiki/Henneguya_salminicola" class="mw-redirect" title="Henneguya salminicola">Henneguya salminicola</a> – was discovered that lacks a mitochondrial genome but retains structures deemed mitochondrion-related organelles. Moreover, nuclear DNA genes involved in aerobic respiration and in mitochondrial DNA replication and transcription were either absent or present only as <a href="/wiki/Pseudogene" title="Pseudogene">pseudogenes</a>. This is the first multicellular organism known to have this absence of aerobic respiration and live completely free of oxygen dependency.<a href="#cite_note-20">[20]</a><a href="#cite_note-21">[21]</a> <div class="mw-heading mw-heading3"><h3 id="Plants_and_fungi">Plants and fungi</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=4" title="Edit section: Plants and fungi">edit</a>]</div> There are three different mitochondrial genome types in plants and fungi. The first type is a circular genome that has introns (type 2) and may range from 19 to 1000 kbp in length. The second genome type is a circular genome (about 20–1000 kbp) that also has a plasmid-like structure (1 kb) (type 3). The final genome type found in plants and fungi is a linear genome made up of homogeneous DNA molecules (type 5).<a href="#cite_note-22">[22]</a><a href="#cite_note-23">[23]</a><a href="#cite_note-24">[24]</a> Great variation in mtDNA gene content and size exists among fungi and plants, although there appears to be a core subset of genes present in all eukaryotes (except for the few that have no mitochondria at all).<a href="#cite_note-Johnston,_I._G._and_Williams,_B._P._2016-9">[9]</a> In Fungi, however, there is no single gene shared among all mitogenomes.<a href="#cite_note-25">[25]</a> Some plant species have enormous mitochondrial genomes, with <a href="/wiki/Silene_conica" title="Silene conica">Silene conica</a> mtDNA containing as many as 11,300,000 base pairs.<a href="#cite_note-26">[26]</a> Surprisingly, even those huge mtDNAs contain the same number and kinds of genes as related plants with much smaller mtDNAs.<a href="#cite_note-27">[27]</a> The genome of the mitochondrion of the cucumber (<a href="/wiki/Cucumis_sativus" class="mw-redirect" title="Cucumis sativus">Cucumis sativus</a>) consists of three circular chromosomes (lengths 1556, 84 and 45 kilobases), which are entirely or largely autonomous with regard to their <a href="/wiki/DNA_replication" title="DNA replication">replication</a>.<a href="#cite_note-Alverson2011-28">[28]</a> <div class="mw-heading mw-heading3"><h3 id="Protists">Protists</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=5" title="Edit section: Protists">edit</a>]</div> <a href="/wiki/Protist" title="Protist">Protists</a> contain the most diverse mitochondrial genomes, with five different types found in this kingdom. Type 2, type 3 and type 5 of the plant and fungal genomes also exist in some protists, as do two unique genome types. One of these unique types is a heterogeneous collection of circular DNA molecules (type 4) while the other is a heterogeneous collection of linear molecules (type 6). Genome types 4 and 6 each range from 1–200 kbp in size.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The smallest mitochondrial genome sequenced to date is the 5,967 bp mtDNA of the parasite <a href="/wiki/Plasmodium_falciparum" title="Plasmodium falciparum">Plasmodium falciparum</a>.<a href="#cite_note-29">[29]</a><a href="#cite_note-30">[30]</a> <a href="/wiki/Endosymbiotic" class="mw-redirect" title="Endosymbiotic">Endosymbiotic</a> gene transfer, the process by which genes that were coded in the mitochondrial genome are transferred to the cell's main genome, likely explains why more complex organisms such as humans have smaller mitochondrial genomes than simpler organisms such as protists.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <table class="wikitable"> <tbody><tr> <th>Genome Type<a href="#cite_note-Kolesnikov,_A._A._2012-14">[14]</a> </th> <th>Kingdom </th> <th>Introns </th> <th>Size </th> <th>Shape </th> <th>Description </th></tr> <tr> <td>1 </td> <td>Animal </td> <td>No </td> <td>11–28 kbp </td> <td>Circular </td> <td>Single molecule </td></tr> <tr> <td>2 </td> <td>Fungi, Plant, Protista </td> <td>Yes </td> <td>19–1000 kbp </td> <td>Circular </td> <td>Single molecule </td></tr> <tr> <td>3 </td> <td>Fungi, Plant, Protista </td> <td>No </td> <td>20–1000 kbp </td> <td>Circular </td> <td>Large molecule and small plasmid like structures </td></tr> <tr> <td>4 </td> <td>Protista </td> <td>No </td> <td>1–200 kbp </td> <td>Circular </td> <td>Heterogeneous group of molecules </td></tr> <tr> <td>5 </td> <td>Fungi, Plant, Protista </td> <td>No </td> <td>1–200 kbp </td> <td>Linear </td> <td>Homogeneous group of molecules </td></tr> <tr> <td>6 </td> <td>Protista </td> <td>No </td> <td>1–200 kbp </td> <td>Linear </td> <td>Heterogeneous group of molecules </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Replication">Replication</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=6" title="Edit section: Replication">edit</a>]</div> Mitochondrial DNA is replicated by the <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a> gamma complex which is composed of a 140 kDa catalytic DNA polymerase encoded by the <a href="/wiki/POLG" title="POLG">POLG</a> gene and two 55 kDa accessory subunits encoded by the <a href="/wiki/POLG2" title="POLG2">POLG2</a> gene.<a href="#cite_note-31">[31]</a> The <a href="/wiki/Replisome" title="Replisome">replisome</a> machinery is formed by DNA polymerase, <a href="/wiki/PEO1" class="mw-redirect" title="PEO1">TWINKLE</a> and mitochondrial <a href="/wiki/SSB_protein" class="mw-redirect" title="SSB protein">SSB proteins</a>. TWINKLE is a <a href="/wiki/Helicase" title="Helicase">helicase</a>, which unwinds short stretches of dsDNA in the 5' to 3' direction.<a href="#cite_note-Jemt2011-32">[32]</a> All these polypeptides are encoded in the nuclear genome.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] During <a href="/wiki/Embryogenesis" class="mw-redirect" title="Embryogenesis">embryogenesis</a>, replication of mtDNA is strictly down-regulated from the fertilized oocyte through the preimplantation embryo.<a href="#cite_note-John2010-33">[33]</a> The resulting reduction in per-cell copy number of mtDNA plays a role in the mitochondrial bottleneck, exploiting <a href="/wiki/Cellular_noise" title="Cellular noise">cell-to-cell variability</a> to ameliorate the inheritance of damaging mutations.<a href="#cite_note-pmid26035426-34">[34]</a> According to Justin St. John and colleagues, "At the <a href="/wiki/Blastocyst" title="Blastocyst">blastocyst</a> stage, the onset of mtDNA replication is specific to the cells of the <a href="/wiki/Trophectoderm" class="mw-redirect" title="Trophectoderm">trophectoderm</a>.<a href="#cite_note-John2010-33">[33]</a> In contrast, the cells of the <a href="/wiki/Inner_cell_mass" title="Inner cell mass">inner cell mass</a> restrict mtDNA replication until they receive the <a href="/wiki/Cell_signaling" title="Cell signaling">signals</a> to <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiate</a> to specific cell types."<a href="#cite_note-John2010-33">[33]</a> <figure class="mw-default-size 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><a href="/wiki/Human_mitochondrial_DNA" class="mw-redirect" title="Human mitochondrial DNA">Human mitochondrial DNA</a> with the 37 genes on their respective H- and L-strands</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Genes_on_the_human_mtDNA_and_their_transcription">Genes on the human mtDNA and their transcription</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=7" title="Edit section: Genes on the human mtDNA and their transcription">edit</a>]</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/Human_mitochondrial_genetics" title="Human mitochondrial genetics">Human mitochondrial genetics</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_karyotype_with_bands_and_sub-bands.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/220px-Human_karyotype_with_bands_and_sub-bands.png" decoding="async" width="220" height="364" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/330px-Human_karyotype_with_bands_and_sub-bands.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/440px-Human_karyotype_with_bands_and_sub-bands.png 2x" data-file-width="9684" data-file-height="16008" /></a><figcaption>Schematic <a href="/wiki/Karyogram" class="mw-redirect" title="Karyogram">karyogram</a> showing the <a href="/wiki/Human_genome" title="Human genome">human genome</a>, with 23 chromosome pairs as well as the mitochondrial genome (to scale at bottom left, annotated "MT"). Its genome is relatively tiny compared to the rest, and its copy number per human cell varies from 0 (<a href="/wiki/Erythrocytes" class="mw-redirect" title="Erythrocytes">erythrocytes</a>)<a href="#cite_note-pmid3178814-35">[35]</a> up to 1,500,000 (<a href="/wiki/Egg_cell" title="Egg cell">oocytes</a>).<a href="#cite_note-pmid28721182-36">[36]</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Karyotype" title="Karyotype">Karyotype</a></div></figcaption></figure> The two strands of the human mitochondrial DNA are distinguished as the heavy strand and the light strand. The heavy strand is rich in <a href="/wiki/Guanine" title="Guanine">guanine</a> and encodes 12 subunits of the <a href="/wiki/Oxidative_phosphorylation" title="Oxidative phosphorylation">oxidative phosphorylation</a> system, two <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNAs</a> (12S and 16S), and 14 <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNAs</a> (tRNAs). The light strand encodes one subunit, and 8 tRNAs. So, altogether mtDNA encodes for two rRNAs, 22 tRNAs, and 13 <a href="/wiki/Protein_subunit" title="Protein subunit">protein subunits</a>, all of which are involved in the oxidative phosphorylation process.<a href="#cite_note-pmid29945721-37">[37]</a><a href="#cite_note-pmid31064115-38">[38]</a> <ul><li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/nuccore/NC_012920.1?report=graph">The complete sequence of the human mitochondrial DNA in graphic form</a></li></ul> <table class="wikitable sortable" style="text-align: center;"> <caption>The 37 genes of the <a href="/wiki/Cambridge_Reference_Sequence" title="Cambridge Reference Sequence">Cambridge Reference Sequence</a> for human mitochondrial DNA and their locations<a href="#cite_note-NCBI_NC_012920-39">[39]</a> </caption> <tbody><tr> <th>Gene </th> <th>Type </th> <th>Product </th> <th>Positions in the mitogenome </th> <th>Strand </th></tr> <tr> <td><a href="/wiki/MT-ATP8" title="MT-ATP8">MT-ATP8</a> </td> <td><a href="/wiki/Protein-coding_gene" class="mw-redirect" title="Protein-coding gene">protein coding</a> </td> <td><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a>, Fo subunit 8 (complex V) </td> <td>08,366–08,572 (overlap with MT-ATP6) </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ATP6" title="MT-ATP6">MT-ATP6</a> </td> <td>protein coding </td> <td><a href="/wiki/ATP_synthase" title="ATP synthase">ATP synthase</a>, Fo subunit 6 (complex V) </td> <td>08,527–09,207 (overlap with MT-ATP8) </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-CO1" class="mw-redirect" title="MT-CO1">MT-CO1</a> </td> <td>protein coding </td> <td><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Cytochrome c oxidase</a>, subunit 1 (complex IV) </td> <td>05,904–07,445 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-CO2" class="mw-redirect" title="MT-CO2">MT-CO2</a> </td> <td>protein coding </td> <td><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Cytochrome c oxidase</a>, subunit 2 (complex IV) </td> <td>07,586–08,269 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-CO3" class="mw-redirect" title="MT-CO3">MT-CO3</a> </td> <td>protein coding </td> <td><a href="/wiki/Cytochrome_c_oxidase" title="Cytochrome c oxidase">Cytochrome c oxidase</a>, subunit 3 (complex IV) </td> <td>09,207–09,990 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-CYB" title="MT-CYB">MT-CYB</a> </td> <td>protein coding </td> <td><a href="/wiki/Cytochrome_b" title="Cytochrome b">Cytochrome b</a> (complex III) </td> <td>14,747–15,887 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND1" title="MT-ND1">MT-ND1</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 1 (complex I) </td> <td>03,307–04,262 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND2" title="MT-ND2">MT-ND2</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 2 (complex I) </td> <td>04,470–05,511 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND3" title="MT-ND3">MT-ND3</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 3 (complex I) </td> <td>10,059–10,404 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND4L" title="MT-ND4L">MT-ND4L</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 4L (complex I) </td> <td>10,470–10,766 (overlap with MT-ND4) </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND4" title="MT-ND4">MT-ND4</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 4 (complex I) </td> <td>10,760–12,137 (overlap with MT-ND4L) </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND5" title="MT-ND5">MT-ND5</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 5 (complex I) </td> <td>12,337–14,148 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-ND6" title="MT-ND6">MT-ND6</a> </td> <td>protein coding </td> <td><a href="/wiki/NADH_dehydrogenase" title="NADH dehydrogenase">NADH dehydrogenase</a>, subunit 6 (complex I) </td> <td>14,149–14,673 </td> <td>L </td></tr> <tr> <td><a href="/wiki/MT-RNR2" title="MT-RNR2">MT-RNR2</a> </td> <td>protein coding </td> <td><a href="/wiki/Humanin" title="Humanin">Humanin</a> </td> <td>— </td> <td>— </td></tr> <tr> <td><a href="/wiki/MT-TA" title="MT-TA">MT-TA</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Alanine" title="Alanine">Alanine</a> (Ala or A) </td> <td>05,587–05,655 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TR" title="MT-TR">MT-TR</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Arginine" title="Arginine">Arginine</a> (Arg or R) </td> <td>10,405–10,469 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TN" title="MT-TN">MT-TN</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Asparagine" title="Asparagine">Asparagine</a> (Asn or N) </td> <td>05,657–05,729 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TD" title="MT-TD">MT-TD</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Aspartic_acid" title="Aspartic acid">Aspartic acid</a> (Asp or D) </td> <td>07,518–07,585 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TC" title="MT-TC">MT-TC</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Cysteine" title="Cysteine">Cysteine</a> (Cys or C) </td> <td>05,761–05,826 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TE" title="MT-TE">MT-TE</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Glutamic_acid" title="Glutamic acid">Glutamic acid</a> (Glu or E) </td> <td>14,674–14,742 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TQ" title="MT-TQ">MT-TQ</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Glutamine" title="Glutamine">Glutamine</a> (Gln or Q) </td> <td>04,329–04,400 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TG" title="MT-TG">MT-TG</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Glycine" title="Glycine">Glycine</a> (Gly or G) </td> <td>09,991–10,058 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TH" title="MT-TH">MT-TH</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Histidine" title="Histidine">Histidine</a> (His or H) </td> <td>12,138–12,206 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TI" title="MT-TI">MT-TI</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Isoleucine" title="Isoleucine">Isoleucine</a> (Ile or I) </td> <td>04,263–04,331 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TL1" title="MT-TL1">MT-TL1</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Leucine" title="Leucine">Leucine</a> (Leu-UUR or L) </td> <td>03,230–03,304 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TL2" title="MT-TL2">MT-TL2</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Leucine" title="Leucine">Leucine</a> (Leu-CUN or L) </td> <td>12,266–12,336 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TK" title="MT-TK">MT-TK</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Lysine" title="Lysine">Lysine</a> (Lys or K) </td> <td>08,295–08,364 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TM" title="MT-TM">MT-TM</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Methionine" title="Methionine">Methionine</a> (Met or M) </td> <td>04,402–04,469 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TF" title="MT-TF">MT-TF</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Phenylalanine" title="Phenylalanine">Phenylalanine</a> (Phe or F) </td> <td>00,577–00,647 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TP" title="MT-TP">MT-TP</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Proline" title="Proline">Proline</a> (Pro or P) </td> <td>15,956–16,023 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TS1" title="MT-TS1">MT-TS1</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Serine" title="Serine">Serine</a> (Ser-UCN or S) </td> <td>07,446–07,514 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TS2" title="MT-TS2">MT-TS2</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Serine" title="Serine">Serine</a> (Ser-AGY or S) </td> <td>12,207–12,265 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TT" title="MT-TT">MT-TT</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Threonine" title="Threonine">Threonine</a> (Thr or T) </td> <td>15,888–15,953 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TW" title="MT-TW">MT-TW</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Tryptophan" title="Tryptophan">Tryptophan</a> (Trp or W) </td> <td>05,512–05,579 </td> <td>H </td></tr> <tr align="center"> <td><a href="/wiki/MT-TY" title="MT-TY">MT-TY</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Tyrosine" title="Tyrosine">Tyrosine</a> (Tyr or Y) </td> <td>05,826–05,891 </td> <td>L </td></tr> <tr align="center"> <td><a href="/wiki/MT-TV_(mitochondrial)" title="MT-TV (mitochondrial)">MT-TV</a> </td> <td>transfer RNA </td> <td>tRNA-<a href="/wiki/Valine" title="Valine">Valine</a> (Val or V) </td> <td>01,602–01,670 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-RNR1" title="MT-RNR1">MT-RNR1</a> </td> <td>ribosomal RNA </td> <td>Small subunit : SSU (12S) </td> <td>00,648–01,601 </td> <td>H </td></tr> <tr> <td><a href="/wiki/MT-RNR2" title="MT-RNR2">MT-RNR2</a> </td> <td>ribosomal RNA </td> <td>Large subunit : LSU (16S) </td> <td>01,671–03,229 </td> <td>H </td></tr></tbody></table> Between most (but not all) protein-coding regions, tRNAs are present (see the <a href="#Map_of_the_human_mitochondrial_genome">human mitochondrial genome map</a>). During transcription, the tRNAs acquire their characteristic L-shape that gets recognized and cleaved by specific enzymes. With the mitochondrial RNA processing, individual mRNA, rRNA, and tRNA sequences are released from the primary transcript.<a href="#cite_note-40">[40]</a> Folded tRNAs therefore act as secondary structure punctuations.<a href="#cite_note-41">[41]</a> <div class="mw-heading mw-heading3"><h3 id="Regulation_of_transcription">Regulation of transcription</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=8" title="Edit section: Regulation of transcription">edit</a>]</div> The promoters for the initiation of the transcription of the heavy and light strands are located in the main non-coding region of the mtDNA called the displacement loop, the <a href="/wiki/D-loop" title="D-loop">D-loop</a>.<a href="#cite_note-pmid29945721-37">[37]</a> There is evidence that the transcription of the mitochondrial rRNAs is regulated by the heavy-strand promoter 1 (HSP1), and the transcription of the polycistronic transcripts coding for the protein subunits are regulated by HSP2.<a href="#cite_note-pmid29945721-37">[37]</a> Measurement of the levels of the mtDNA-encoded RNAs in bovine tissues has shown that there are major differences in the expression of the mitochondrial RNAs relative to total tissue RNA.<a href="#cite_note-1993-Raikhinstein-42">[42]</a> Among the 12 tissues examined the highest level of expression was observed in heart, followed by brain and steroidogenic tissue samples.<a href="#cite_note-1993-Raikhinstein-42">[42]</a> As demonstrated by the effect of the trophic hormone <a href="/wiki/ACTH" class="mw-redirect" title="ACTH">ACTH</a> on adrenal cortex cells, the expression of the mitochondrial genes may be strongly regulated by external factors, apparently to enhance the synthesis of mitochondrial proteins necessary for energy production.<a href="#cite_note-1993-Raikhinstein-42">[42]</a> Interestingly, while the expression of protein-encoding genes was stimulated by ACTH, the levels of the mitochondrial 16S rRNA showed no significant change.<a href="#cite_note-1993-Raikhinstein-42">[42]</a> <div class="mw-heading mw-heading2"><h2 id="Mitochondrial_inheritance">Mitochondrial inheritance</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=9" title="Edit section: Mitochondrial inheritance">edit</a>]</div> In most <a href="/wiki/Metazoans" class="mw-redirect" title="Metazoans">multicellular organisms</a>, mtDNA is inherited from the mother (maternally inherited). Mechanisms for this include simple dilution (an egg contains on average 200,000 mtDNA molecules, whereas a healthy human <a href="/wiki/Sperm" title="Sperm">sperm</a> has been reported to contain on average 5 molecules),<a href="#cite_note-43">[43]</a><a href="#cite_note-44">[44]</a> degradation of sperm mtDNA in the male genital tract and in the fertilized egg; and, at least in a few organisms, failure of sperm mtDNA to enter the egg. Whatever the mechanism, this single parent (<a href="/wiki/Uniparental_inheritance" title="Uniparental inheritance">uniparental inheritance</a>) pattern of mtDNA inheritance is found in most animals, most plants and also in fungi.<a href="#cite_note-45">[45]</a> In a study published in 2018, human babies were reported to inherit mtDNA from both their fathers and their mothers resulting in mtDNA <a href="/wiki/Heteroplasmy" title="Heteroplasmy">heteroplasmy</a>,<a href="#cite_note-Biparental_Inheritance_of_Mitochond-46">[46]</a> a finding that has been rejected by other scientists.<a href="#cite_note-pmid34031572-47">[47]</a><a href="#cite_note-pmid32330850-48">[48]</a><a href="#cite_note-pmid32269217-49">[49]</a> <div class="mw-heading mw-heading3"><h3 id="Female_inheritance">Female inheritance</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=10" title="Edit section: Female inheritance">edit</a>]</div> In <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a>, mitochondria are normally inherited exclusively from the mother; the mitochondria in mammalian sperm are usually destroyed by the egg cell after fertilization. Also, mitochondria are present solely in the midpiece, which is used for propelling the sperm cells, and sometimes the midpiece, along with the tail, is lost during fertilization. In 1999 it was reported that paternal sperm mitochondria (containing mtDNA) 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-50">[50]</a> Some <a href="/wiki/In_vitro" title="In vitro">in vitro</a> fertilization techniques, particularly injecting a sperm into an <a href="/wiki/Oocyte" title="Oocyte">oocyte</a>, may interfere with this.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The fact that mitochondrial DNA is mostly maternally inherited enables <a href="/wiki/Genetic_genealogy" title="Genetic genealogy">genealogical</a> researchers to trace <a href="/wiki/Matrilineality" title="Matrilineality">maternal lineage</a> far back in time. (<a href="/wiki/Y_chromosome" title="Y chromosome">Y-chromosomal DNA</a>, paternally inherited, is used in an analogous way to determine the <a href="/wiki/Patrilineal" class="mw-redirect" title="Patrilineal">patrilineal</a> history.) This is usually accomplished on <a href="/wiki/Human_mitochondrial_DNA" class="mw-redirect" title="Human mitochondrial DNA">human mitochondrial DNA</a> by sequencing the <a href="/wiki/Hypervariable_control_region" class="mw-redirect" title="Hypervariable control region">hypervariable control regions</a> (HVR1 or HVR2), and sometimes the complete molecule of the mitochondrial DNA, as a <a href="/wiki/Genealogical_DNA_test" title="Genealogical DNA test">genealogical DNA test</a>.<a href="#cite_note-51">[51]</a> HVR1, for example, consists of about 440 base pairs. These 440 base pairs are compared to the same regions of other individuals (either specific people or subjects in a database) to determine maternal lineage. Most often, the comparison is made with the revised <a href="/wiki/Cambridge_Reference_Sequence" title="Cambridge Reference Sequence">Cambridge Reference Sequence</a>. Vilà et al. have published studies tracing the matrilineal descent of domestic dogs from wolves.<a href="#cite_note-Vila-52">[52]</a> The concept of the <a href="/wiki/Mitochondrial_Eve" title="Mitochondrial Eve">Mitochondrial Eve</a> is based on the same type of analysis, attempting to discover the origin of <a href="/wiki/Human" title="Human">humanity</a> by tracking the lineage back in time.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="The_mitochondrial_bottleneck">The mitochondrial bottleneck</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=11" title="Edit section: The mitochondrial bottleneck">edit</a>]</div> Entities subject to 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 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">random 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 in which different cells have different mutant loads. Cell-level selection may then act to remove those cells with more mutant mtDNA, leading to a stabilisation or reduction in mutant load between generations. The mechanism underlying the bottleneck is debated,<a href="#cite_note-53">[53]</a><a href="#cite_note-54">[54]</a><a href="#cite_note-55">[55]</a><a href="#cite_note-56">[56]</a> with a recent mathematical and experimental metastudy providing evidence for a combination of the random partitioning of mtDNAs at cell divisions and the random turnover of mtDNA molecules within the cell.<a href="#cite_note-pmid26035426-34">[34]</a> <div class="mw-heading mw-heading3"><h3 id="Male_inheritance">Male inheritance</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=12" title="Edit section: Male inheritance">edit</a>]</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/Paternal_mtDNA_transmission" title="Paternal mtDNA transmission">Paternal mtDNA transmission</a></div> Male mitochondrial DNA inheritance has been discovered in <a href="/wiki/Plymouth_Rock_chicken" title="Plymouth Rock chicken">Plymouth Rock chickens</a>.<a href="#cite_note-57">[57]</a> Evidence supports rare instances of male mitochondrial inheritance in some mammals as well. Specifically, documented occurrences exist for mice,<a href="#cite_note-58">[58]</a><a href="#cite_note-59">[59]</a> where the male-inherited mitochondria were subsequently rejected. It has also been found in sheep,<a href="#cite_note-60">[60]</a> and in cloned cattle.<a href="#cite_note-61">[61]</a> Rare cases of male mitochondrial inheritance have been documented in humans.<a href="#cite_note-62">[62]</a><a href="#cite_note-63">[63]</a><a href="#cite_note-64">[64]</a><a href="#cite_note-Biparental_Inheritance_of_Mitochond-46">[46]</a> Although many of these cases involve cloned embryos or subsequent rejection of the paternal mitochondria, others document <a href="/wiki/In_vivo" title="In vivo">in vivo</a> inheritance and persistence under lab conditions.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Doubly uniparental inheritance of mtDNA is observed in bivalve mollusks. In those species, females have only one type of mtDNA (F), whereas males have F type mtDNA in their somatic cells, but M type of mtDNA (which can be as much as 30% divergent) in <a href="/wiki/Germline" title="Germline">germline</a> cells.<a href="#cite_note-65">[65]</a> Paternally inherited mitochondria have additionally been reported in some insects such as <a href="/wiki/Drosophila" title="Drosophila">fruit flies</a>,<a href="#cite_note-66">[66]</a><a href="#cite_note-67">[67]</a> <a href="/wiki/Honeybee" class="mw-redirect" title="Honeybee">honeybees</a>,<a href="#cite_note-68">[68]</a> and <a href="/wiki/Magicicada" class="mw-redirect" title="Magicicada">periodical cicadas</a>.<a href="#cite_note-69">[69]</a> <div class="mw-heading mw-heading3"><h3 id="Mitochondrial_donation">Mitochondrial donation</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=13" title="Edit section: Mitochondrial donation">edit</a>]</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_donation" class="mw-redirect" title="Mitochondrial donation">Mitochondrial donation</a></div> An IVF technique known as mitochondrial donation or mitochondrial replacement therapy (MRT) results in offspring containing mtDNA from a donor female, and nuclear DNA from the mother and father. In the spindle transfer procedure, the nucleus of an egg is inserted into the cytoplasm of an egg from a donor female which has had its nucleus removed, but still contains the donor female's mtDNA. The composite egg is then fertilized with the male's sperm. The procedure is used when a woman with <a href="/wiki/Mitochondrial_disease" title="Mitochondrial disease">genetically defective mitochondria</a> wishes to procreate and produce offspring with healthy mitochondria.<a href="#cite_note-Frith-70">[70]</a> The first known child to be born as a result of mitochondrial donation was a boy born to a Jordanian couple in Mexico on 6 April 2016.<a href="#cite_note-71">[71]</a> <div class="mw-heading mw-heading2"><h2 id="Mutations_and_disease">Mutations and disease</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=14" title="Edit section: Mutations and disease">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Mitochondrial_DNA_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Mitochondrial_DNA_en.svg/300px-Mitochondrial_DNA_en.svg.png" decoding="async" width="300" height="242" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Mitochondrial_DNA_en.svg/450px-Mitochondrial_DNA_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Mitochondrial_DNA_en.svg/600px-Mitochondrial_DNA_en.svg.png 2x" data-file-width="1930" data-file-height="1560" /></a><figcaption><a href="/wiki/Human_mitochondrial_DNA" class="mw-redirect" title="Human mitochondrial DNA">Human mitochondrial DNA</a> with groups of protein-, rRNA- and tRNA-encoding genes</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Mitochondrial_DNA_and_diseases.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mitochondrial_DNA_and_diseases.svg/300px-Mitochondrial_DNA_and_diseases.svg.png" decoding="async" width="300" height="242" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mitochondrial_DNA_and_diseases.svg/450px-Mitochondrial_DNA_and_diseases.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mitochondrial_DNA_and_diseases.svg/600px-Mitochondrial_DNA_and_diseases.svg.png 2x" data-file-width="1734" data-file-height="1401" /></a><figcaption>The involvement of mitochondrial DNA in several human diseases</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Susceptibility">Susceptibility</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=15" title="Edit section: Susceptibility">edit</a>]</div> The concept that mtDNA is particularly susceptible to <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> generated by the <a href="/wiki/Respiratory_chain" class="mw-redirect" title="Respiratory chain">respiratory chain</a> due to its proximity remains controversial.<a href="#cite_note-pmid19796285-72">[72]</a> mtDNA does not accumulate any more oxidative base damage than nuclear DNA.<a href="#cite_note-73">[73]</a> It has been reported that at least some types of oxidative DNA damage are repaired more efficiently in mitochondria than they are in the nucleus.<a href="#cite_note-74">[74]</a> mtDNA is packaged with proteins which appear to be as protective as proteins of the nuclear chromatin.<a href="#cite_note-75">[75]</a> Moreover, mitochondria evolved a unique mechanism which maintains mtDNA integrity through degradation of excessively damaged genomes followed by replication of intact/repaired mtDNA. This mechanism is not present in the nucleus and is enabled by multiple copies of mtDNA present in mitochondria.<a href="#cite_note-76">[76]</a> The outcome of mutation in mtDNA may be an alteration in the coding instructions for some proteins,<a href="#cite_note-77">[77]</a> which may have an effect on organism metabolism and/or fitness. <div class="mw-heading mw-heading3"><h3 id="Genetic_illness">Genetic illness</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=16" title="Edit section: Genetic illness">edit</a>]</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/Mitochondrial_disease" title="Mitochondrial disease">Mitochondrial disease</a></div> Mutations of mitochondrial DNA can lead to a number of illnesses including <a href="/wiki/Exercise_intolerance" title="Exercise intolerance">exercise intolerance</a> and <a href="/wiki/Kearns%E2%80%93Sayre_syndrome" title="Kearns–Sayre syndrome">Kearns–Sayre syndrome</a> (KSS), which causes a person to lose full function of heart, eye, and muscle movements. Some evidence suggests that they might be major contributors to the aging process and <a href="/wiki/Aging-associated_diseases" title="Aging-associated diseases">age-associated pathologies</a>.<a href="#cite_note-78">[78]</a> Particularly in the context of disease, the proportion of mutant mtDNA molecules in a cell is termed <a href="/wiki/Heteroplasmy" title="Heteroplasmy">heteroplasmy</a>. The within-cell and between-cell distributions of heteroplasmy dictate the onset and severity of disease<a href="#cite_note-79">[79]</a> and are influenced by complicated <a href="/wiki/Cellular_noise" title="Cellular noise">stochastic processes</a> within the cell and during development.<a href="#cite_note-pmid26035426-34">[34]</a><a href="#cite_note-80">[80]</a> Mutations in mitochondrial tRNAs can be responsible for severe diseases like the <a href="/wiki/MELAS_syndrome" title="MELAS syndrome">MELAS</a> and <a href="/wiki/MERRF_syndrome" title="MERRF syndrome">MERRF</a> syndromes.<a href="#cite_note-nature-81">[81]</a> Mutations in nuclear genes that encode proteins that mitochondria use can also contribute to mitochondrial diseases. These diseases do not follow mitochondrial inheritance patterns, but instead follow Mendelian inheritance patterns.<a href="#cite_note-82">[82]</a> <div class="mw-heading mw-heading3"><h3 id="Use_in_disease_diagnosis">Use in disease diagnosis</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=17" title="Edit section: Use in disease diagnosis">edit</a>]</div> Recently a mutation in mtDNA has been used to help diagnose prostate cancer in patients with negative <a href="/wiki/Prostate_biopsy" title="Prostate biopsy">prostate biopsy</a>.<a href="#cite_note-pmid20944788-83">[83]</a><a href="#cite_note-pmid20084081-84">[84]</a> mtDNA alterations can be detected in the bio-fluids of patients with cancer.<a href="#cite_note-Mair-85">[85]</a> mtDNA is characterized by the high rate of polymorphisms and mutations. Some of which are increasingly recognized as an important cause of human pathology such as oxidative phosphorylation (OXPHOS) disorders, maternally inherited diabetes and deafness (MIDD), Type 2 diabetes mellitus, <a href="/wiki/Neurodegenerative_disease" title="Neurodegenerative disease">Neurodegenerative disease</a>, heart failure and cancer.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Relationship_with_ageing">Relationship with ageing</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=18" title="Edit section: Relationship with ageing">edit</a>]</div> Though the idea is controversial, some evidence suggests a link between aging and mitochondrial genome dysfunction.<a href="#cite_note-86">[86]</a> In essence, mutations in mtDNA upset a careful balance of <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> (ROS) production and enzymatic ROS scavenging (by enzymes like <a href="/wiki/Superoxide_dismutase" title="Superoxide dismutase">superoxide dismutase</a>, <a href="/wiki/Catalase" title="Catalase">catalase</a>, <a href="/wiki/Glutathione_peroxidase" title="Glutathione peroxidase">glutathione peroxidase</a> and others). However, some mutations that increase ROS production (e.g., by reducing antioxidant defenses) in worms increase, rather than decrease, their longevity.<a href="#cite_note-pmid19796285-72">[72]</a> Also, <a href="/wiki/Naked_mole-rat" title="Naked mole-rat">naked mole rats</a>, <a href="/wiki/Rodent" title="Rodent">rodents</a> about the size of <a href="/wiki/Mouse" title="Mouse">mice</a>, live about eight times longer than mice despite having reduced, compared to mice, antioxidant defenses and increased oxidative damage to biomolecules.<a href="#cite_note-87">[87]</a> Once, there was thought to be a positive feedback loop at work (a 'Vicious Cycle'); as mitochondrial DNA accumulates genetic damage caused by free radicals, the mitochondria lose function and leak free radicals into the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>. A decrease in mitochondrial function reduces overall metabolic efficiency.<a href="#cite_note-88">[88]</a> However, this concept was conclusively disproved when it was demonstrated that mice, which were genetically altered to accumulate mtDNA mutations at accelerated rate do age prematurely, but their tissues do not produce more ROS as predicted by the 'Vicious Cycle' hypothesis.<a href="#cite_note-89">[89]</a> Supporting a link between longevity and mitochondrial DNA, some studies have found correlations between biochemical properties of the mitochondrial DNA and the longevity of species.<a href="#cite_note-90">[90]</a> The application of a mitochondrial-specific ROS scavenger, which lead to a significant longevity of the mice studied,<a href="#cite_note-91">[91]</a> suggests that mitochondria may still be well-implicated in ageing. Extensive research is being conducted to further investigate this link and methods to combat ageing. Presently, <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a> and <a href="/wiki/Nutraceutical" title="Nutraceutical">nutraceutical</a> supplementation are popular areas of ongoing research.<a href="#cite_note-92">[92]</a><a href="#cite_note-93">[93]</a> Bjelakovic et al. analyzed the results of 78 studies between 1977 and 2012, involving a total of 296,707 participants, and concluded that antioxidant supplements do not reduce all-cause mortality nor extend lifespan, while some of them, such as beta carotene, vitamin E, and higher doses of vitamin A, may actually increase mortality.<a href="#cite_note-94">[94]</a> In a recent study, it was shown that dietary restriction can reverse ageing alterations by affecting the accumulation of mtDNA damage in several organs of rats. For example, dietary restriction prevented age-related accumulation of mtDNA damage in the cortex and decreased it in the lung and testis.<a href="#cite_note-95">[95]</a> <div class="mw-heading mw-heading3"><h3 id="Neurodegenerative_diseases">Neurodegenerative diseases</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=19" title="Edit section: Neurodegenerative diseases">edit</a>]</div> Increased mt<a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage</a> is a feature of several <a href="/wiki/Neurodegeneration" class="mw-redirect" title="Neurodegeneration">neurodegenerative diseases</a>. The brains of individuals with <a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a> have elevated levels of <a href="/wiki/DNA_oxidation" title="DNA oxidation">oxidative DNA damage</a> in both <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a> and mtDNA, but the mtDNA has approximately 10-fold higher levels than nuclear DNA.<a href="#cite_note-pmid15857398-96">[96]</a> It has been proposed that aged <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> is the critical factor in the origin of neurodegeneration in Alzheimer's disease.<a href="#cite_note-pmid24733654-97">[97]</a> Analysis of the brains of AD patients suggested an impaired function of the <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> pathway, which would cause reduce the overall quality of mtDNA.<a href="#cite_note-98">[98]</a> In <a href="/wiki/Huntington%27s_disease" title="Huntington's disease">Huntington's disease</a>, mutant <a href="/wiki/Huntingtin_protein" class="mw-redirect" title="Huntingtin protein">huntingtin protein</a> causes <a href="/wiki/Mitochondrial_dysfunction" class="mw-redirect" title="Mitochondrial dysfunction">mitochondrial dysfunction</a> involving inhibition of <a href="/wiki/Mitochondrial" class="mw-redirect" title="Mitochondrial">mitochondrial</a> <a href="/wiki/Electron_transport_chain" title="Electron transport chain">electron transport</a>, higher levels of <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> and increased <a href="/wiki/Oxidative_stress" title="Oxidative stress">oxidative stress</a>.<a href="#cite_note-pmid28785371-99">[99]</a> Mutant huntingtin protein promotes oxidative damage to mtDNA, as well as nuclear DNA, that may contribute to Huntington's disease <a href="/wiki/Pathology" title="Pathology">pathology</a>.<a href="#cite_note-pmid23602907-100">[100]</a> The <a href="/wiki/DNA_oxidation" title="DNA oxidation">DNA oxidation</a> product <a href="/wiki/8-oxoguanine" class="mw-redirect" title="8-oxoguanine">8-oxoguanine</a> (8-oxoG) is a well-established marker of oxidative DNA damage. In persons with <a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">amyotrophic lateral sclerosis</a> (ALS), the enzymes that normally repair 8-oxoG DNA damages in the mtDNA of spinal <a href="/wiki/Motor_neuron" title="Motor neuron">motor neurons</a> are impaired.<a href="#cite_note-pmid11904761-101">[101]</a> Thus oxidative damage to mtDNA of motor neurons may be a significant factor in the <a href="/wiki/Etiology" title="Etiology">etiology</a> of ALS.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Correlation_of_the_mtDNA_base_composition_with_animal_life_spans">Correlation of the mtDNA base composition with animal life spans</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=20" title="Edit section: Correlation of the mtDNA base composition with animal life spans">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png/220px-Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png" decoding="async" width="220" height="109" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png/330px-Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png/440px-Correlation_between_the_mtDNA_GC%25_and_maximum_life_span_across_387_different_mammalian_species.png 2x" data-file-width="1682" data-file-height="837" /></a><figcaption>Animal species mtDNA base composition was retrieved from the MitoAge database and compared to their maximum life span from AnAge database.</figcaption></figure> Over the past decade, an Israeli research group led by Professor Vadim Fraifeld has shown that strong and significant <a href="/wiki/Correlation_and_dependence" class="mw-redirect" title="Correlation and dependence">correlations</a> exist between the mtDNA base composition and animal species-specific maximum life spans.<a href="#cite_note-Lehmann_et_al.,_2006-102">[102]</a><a href="#cite_note-Lehmann_et_al.,_2008-103">[103]</a><a href="#cite_note-Lehmann_et_al.,_2013-104">[104]</a> As demonstrated in their work, higher mtDNA <a href="/wiki/Guanine" title="Guanine">guanine</a> + <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> content (<a href="/wiki/GC-content" title="GC-content">GC%</a>) strongly associates with longer <a href="/wiki/Maximum_life_span" title="Maximum life span">maximum life spans</a> across animal species. An additional observation is that the mtDNA GC% correlation with the maximum life spans is independent of the well-known correlation between animal species metabolic rate and maximum life spans. The mtDNA GC% and resting metabolic rate explain the differences in animal species maximum life spans in a multiplicative manner (i.e., species maximum life span = their mtDNA GC% * metabolic rate).<a href="#cite_note-Lehmann_et_al.,_2008-103">[103]</a> To support the scientific community in carrying out comparative analyses between mtDNA features and longevity across animals, a dedicated database was built named <a rel="nofollow" class="external text" href="http://www.mitoage.info/">MitoAge</a>.<a href="#cite_note-105">[105]</a> <div class="mw-heading mw-heading3"><h3 id="mtDNA_mutational_spectrum_is_sensitive_to_species-specific_life-history_traits">mtDNA mutational spectrum is sensitive to species-specific life-history traits</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=21" title="Edit section: mtDNA mutational spectrum is sensitive to species-specific life-history traits">edit</a>]</div> De novo mutations arise either due to mistakes during DNA replication or due to unrepaired damage caused in turn by endogenous and exogenous mutagens. It has been long believed that mtDNA can be particularly sensitive to damage caused by reactive oxygen species (ROS), however G>T substitutions, the hallmark of the oxidative damage in the nuclear genome, are very rare in mtDNA and do not increase with age. Comparing the mtDNA mutational spectra of hundreds of mammalian species, it has been recently demonstrated that species with extended lifespans have an increased rate of A>G substitutions on single-stranded heavy chain.<a href="#cite_note-106">[106]</a> This discovery led to the hypothesis that A>G is a mitochondria-specific marker of age-associated oxidative damage. This finding provides a mutational (contrary to the selective one) explanation for the observation that long-lived species have GC-rich mtDNA: long-lived species become GC-rich simply because of their biased process of mutagenesis. An association between mtDNA mutational spectrum and species-specific life-history traits in mammals opens a possibility to link these factors together discovering new life-history-specific mutagens in different groups of organisms.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Relationship_with_non-B_(non-canonical)_DNA_structures">Relationship with non-B (non-canonical) DNA structures</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=22" title="Edit section: Relationship with non-B (non-canonical) DNA structures">edit</a>]</div> Deletion breakpoints frequently occur within or near regions showing non-canonical (non-B) conformations, namely hairpins, cruciforms and cloverleaf-like elements.<a href="#cite_note-107">[107]</a> Moreover, there is data supporting the involvement of helix-distorting intrinsically curved regions and long G-tetrads in eliciting instability events. In addition, higher breakpoint densities were consistently observed within GC-skewed regions and in the close vicinity of the degenerate sequence motif YMMYMNNMMHM.<a href="#cite_note-108">[108]</a> <div class="mw-heading mw-heading2"><h2 id="Use_in_forensics">Use in forensics</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=23" title="Edit section: Use in forensics">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For use in human identification, see <a href="/wiki/Human_mitochondrial_DNA" class="mw-redirect" title="Human mitochondrial DNA">Human mitochondrial DNA</a>.</div> Unlike nuclear DNA, which is inherited from both parents and in which genes are rearranged in the process of <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a>, there is usually no change in mtDNA from parent to offspring. Although mtDNA also recombines, it does so with copies of itself within the same mitochondrion. Because of this and because the <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> of animal mtDNA is higher than that of nuclear DNA,<a href="#cite_note-Brown-109">[109]</a> mtDNA is a powerful tool for tracking ancestry through females (<a href="/wiki/Matrilineage" class="mw-redirect" title="Matrilineage">matrilineage</a>) and has been used in this role to track the ancestry of many species back hundreds of generations.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] mtDNA testing can be used by forensic scientists in cases where nuclear DNA is severely degraded. Autosomal cells only have two copies of nuclear DNA, but can have hundreds of copies of mtDNA due to the multiple mitochondria present in each cell. This means highly degraded evidence that would not be beneficial for STR analysis could be used in mtDNA analysis. mtDNA may be present in bones, teeth, or hair, which could be the only remains left in the case of severe degradation. In contrast to STR analysis, mtDNA sequencing uses <a href="/wiki/Sanger_sequencing" title="Sanger sequencing">Sanger sequencing</a>. The known sequence and questioned sequence are both compared to the Revised Cambridge Reference Sequence to generate their respective haplotypes. If the known sample sequence and questioned sequence originated from the same matriline, one would expect to see identical sequences and identical differences from the rCRS.<a href="#cite_note-110">[110]</a> Cases arise where there are no known samples to collect and the unknown sequence can be searched in a database such as EMPOP. The Scientific Working Group on DNA Analysis Methods recommends three conclusions for describing the differences between a known mtDNA sequence and a questioned mtDNA sequence: exclusion for two or more differences between the sequences, inconclusive if there is one nucleotide difference, or cannot exclude if there are no nucleotide differences between the two sequences.<a href="#cite_note-111">[111]</a> The rapid mutation rate (in animals) makes mtDNA useful for assessing genetic relationships of individuals or groups within a species and also for identifying and quantifying the phylogeny (evolutionary relationships; see <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogenetics</a>) among different species. To do this, biologists determine and then compare the mtDNA sequences from different individuals or species. Data from the comparisons is used to construct a network of relationships among the sequences, which provides an estimate of the relationships among the individuals or species from which the mtDNAs were taken. mtDNA can be used to estimate the relationship between both closely related and distantly related species. Due to the high mutation rate of mtDNA in animals, the 3rd positions of the codons change relatively rapidly, and thus provide information about the genetic distances among closely related individuals or species. On the other hand, the substitution rate of mt-proteins is very low, thus amino acid changes accumulate slowly (with corresponding slow changes at 1st and 2nd codon positions) and thus they provide information about the genetic distances of distantly related species. Statistical models that treat substitution rates among codon positions separately, can thus be used to simultaneously estimate phylogenies that contain both closely and distantly related species<a href="#cite_note-nature-81">[81]</a> Mitochondrial DNA was admitted into evidence for the first time ever in a United States courtroom in 1996 during State of Tennessee v. Paul Ware.<a href="#cite_note-112">[112]</a> In the 1998 United States court case of Commonwealth of Pennsylvania v. Patricia Lynne Rorrer,<a href="#cite_note-113">[113]</a> mitochondrial DNA was admitted into evidence in the State of Pennsylvania for the first time.<a href="#cite_note-114">[114]</a><a href="#cite_note-115">[115]</a> The case was featured in episode 55 of season 5 of the true crime drama series <a href="/wiki/Forensic_Files_(season_5)" class="mw-redirect" title="Forensic Files (season 5)">Forensic Files (season 5)</a>.<a href="#cite_note-116">[116]</a> Mitochondrial DNA was first admitted into evidence in <a href="/wiki/California" title="California">California</a>, United States, in the successful prosecution of David Westerfield for the 2002 kidnapping and murder of 7-year-old <a href="/wiki/Murder_of_Danielle_van_Dam" title="Murder of Danielle van Dam">Danielle van Dam</a> in <a href="/wiki/San_Diego" title="San Diego">San Diego</a>: it was used for both human and dog identification.<a href="#cite_note-117">[117]</a> This was the first trial in the U.S. to admit canine DNA.<a href="#cite_note-118">[118]</a> The remains of <a href="/wiki/Richard_III_of_England" title="Richard III of England">King Richard III</a>, who died in 1485, were identified by comparing his mtDNA with that of two matrilineal descendants of his sister who were alive in 2013, 527 years after he died.<a href="#cite_note-119">[119]</a> <div class="mw-heading mw-heading2"><h2 id="Use_in_evolutionary_biology_and_systematic_biology">Use in evolutionary biology and systematic biology</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=24" title="Edit section: Use in evolutionary biology and systematic biology">edit</a>]</div> mtDNA is conserved across eukaryotic organism given the critical role of mitochondria in <a href="/wiki/Cellular_respiration" title="Cellular respiration">cellular respiration</a>. However, due to less efficient DNA repair (compared to nuclear DNA) it has a relatively high mutation rate (but slow compared to other DNA regions such as <a href="/wiki/Microsatellite" title="Microsatellite">microsatellites</a>) which makes it useful for studying the evolutionary relationships—<a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogeny</a>—of organisms. Biologists can determine and then compare mtDNA sequences among different species and use the comparisons to build an <a href="/wiki/Evolutionary_tree" class="mw-redirect" title="Evolutionary tree">evolutionary tree</a> for the species examined.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] For instance, while most <a href="/wiki/Nuclear_gene" title="Nuclear gene">nuclear genes</a> are nearly identical between humans and <a href="/wiki/Chimpanzee" title="Chimpanzee">chimpanzees</a>, their mitochondrial genomes are 9.8% different. Human and <a href="/wiki/Western_gorilla" title="Western gorilla">gorilla</a> mitochondrial genomes are 11.8% different, suggesting that humans may be more closely related to chimpanzees than gorillas.<a href="#cite_note-120">[120]</a> <div class="mw-heading mw-heading2"><h2 id="mtDNA_in_nuclear_DNA">mtDNA in nuclear DNA</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=25" title="Edit section: mtDNA in nuclear DNA">edit</a>]</div> <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">Whole genome sequences</a> of more than 66,000 people revealed that most of them had some mitochondrial DNA inserted into their <a href="/wiki/Cell_nucleus" title="Cell nucleus">nuclear</a> <a href="/wiki/Genome" title="Genome">genomes</a>. More than 90% of these nuclear-mitochondrial segments (<a href="/wiki/NUMT" class="mw-redirect" title="NUMT">NUMTs</a>) were inserted after humans diverged from the other <a href="/wiki/Ape" title="Ape">apes</a>. Results indicate such transfers currently occur as frequent as once in every ≈4,000 human births.<a href="#cite_note-121">[121]</a> It appears that <a href="/wiki/Organelle" title="Organelle">organellar</a> DNA is much more often transferred to nuclear DNA than previously thought. This observation also supports the idea of the <a href="/wiki/Symbiogenesis" title="Symbiogenesis">endosymbiont theory</a> that <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> have evolved from <a href="/wiki/Endosymbiont" title="Endosymbiont">endosymbionts</a> which turned into organelles while transferring most of their DNA to the nucleus so that the organellar genome shrunk in the process.<a href="#cite_note-122">[122]</a> <div class="mw-heading mw-heading2"><h2 id="History">History</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=26" title="Edit section: History">edit</a>]</div> Mitochondrial DNA was discovered in the 1960s by Margit M. K. Nass and Sylvan Nass by <a href="/wiki/Electron_microscope" title="Electron microscope">electron microscopy</a> as DNase-sensitive threads inside mitochondria,<a href="#cite_note-123">[123]</a> and by Ellen Haslbrunner, <a href="/wiki/Hans_Tuppy" title="Hans Tuppy">Hans Tuppy</a> and <a href="/wiki/Gottfried_Schatz" title="Gottfried Schatz">Gottfried Schatz</a> by biochemical assays on highly purified mitochondrial fractions.<a href="#cite_note-124">[124]</a> <div class="mw-heading mw-heading2"><h2 id="Mitochondrial_sequence_databases">Mitochondrial sequence databases</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=27" title="Edit section: Mitochondrial sequence databases">edit</a>]</div> Several specialized databases have been founded to collect mitochondrial genome sequences and other information. Although most of them focus on sequence data, some of them include phylogenetic or functional information. <ul><li>AmtDB: a database of ancient human mitochondrial genomes.<a href="#cite_note-AmtDB-125">[125]</a></li> <li>InterMitoBase: an annotated database and analysis platform of protein-protein interactions for human mitochondria.<a href="#cite_note-InterMitoBase-126">[126]</a> (apparently last updated in 2010, but still available)</li> <li>MitoBreak: the mitochondrial DNA breakpoints database.<a href="#cite_note-MitoBreak-127">[127]</a></li> <li>MitoFish and MitoAnnotator: a mitochondrial genome database of fish.<a href="#cite_note-MitoFish-128">[128]</a> See also Cawthorn et al.<a href="#cite_note-Cawthorn-129">[129]</a></li> <li>Mitome: a database for comparative mitochondrial genomics in metazoan animals<a href="#cite_note-Mitome-130">[130]</a> (no longer available)</li> <li>MitoRes: a resource of nuclear-encoded mitochondrial genes and their products in metazoa<a href="#cite_note-MitoRes-131">[131]</a> (apparently no longer being updated)</li> <li>MitoSatPlant: Mitochondrial microsatellites database of viridiplantae.<a href="#cite_note-MitoSatPlant-132">[132]</a></li> <li>MitoZoa 2.0: a database for comparative and evolutionary analyses of mitochondrial genomes in Metazoa.<a href="#cite_note-MitoZoa-133">[133]</a> (no longer available)</li></ul> <div class="mw-heading mw-heading2"><h2 id="MtDNA-phenotype_association_databases">MtDNA-phenotype association databases</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=28" title="Edit section: MtDNA-phenotype association databases">edit</a>]</div> <a href="/wiki/Genome-wide_association_study" title="Genome-wide association study">Genome-wide association studies</a> can reveal associations of mtDNA genes and their mutations with <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a> including <a href="/wiki/Longevity" title="Longevity">lifespan</a> and disease risks. In 2021, the largest, <a href="/wiki/UK_Biobank" title="UK Biobank">UK Biobank</a>-based, genome-wide association study of mitochondrial DNA unveiled 260 new associations with phenotypes including <a href="/wiki/Longevity" title="Longevity">lifespan</a> and disease risks for e.g. type 2 diabetes.<a href="#cite_note-134">[134]</a><a href="#cite_note-135">[135]</a> <div class="mw-heading mw-heading3"><h3 id="Mitochondrial_mutation_databases">Mitochondrial mutation databases</h3>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=29" title="Edit section: Mitochondrial mutation databases">edit</a>]</div> Several specialized databases exist that report polymorphisms and mutations in the human mitochondrial DNA, together with the assessment of their pathogenicity. <ul><li>MitImpact: A collection of pre-computed pathogenicity predictions for all nucleotide changes that cause non-synonymous substitutions in human mitochondrial protein coding genes <a rel="nofollow" class="external text" href="http://mitimpact.css-mendel.it/">MitImpact 3D - IRCCS-CSS Bioinformatics lab</a>.</li> <li>MITOMAP: A compendium of polymorphisms and mutations in human mitochondrial DNA <a rel="nofollow" class="external text" href="http://www.mitomap.org/MITOMAP">WebHome < MITOMAP < Foswiki</a>.</li></ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=30" title="Edit section: See also">edit</a>]</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: 30em;"> <ul><li><a href="/wiki/Clade" title="Clade">Clade</a></li> <li><a href="/wiki/CoRR_hypothesis" title="CoRR hypothesis">CoRR hypothesis</a></li> <li><a href="/wiki/Haplogroup" title="Haplogroup">Haplogroup</a></li> <li><a href="/wiki/Heteroplasmy" title="Heteroplasmy">Heteroplasmy</a></li> <li><a href="/wiki/Human_mitochondrial_DNA_haplogroup" title="Human mitochondrial DNA haplogroup">Human mitochondrial DNA haplogroup</a></li> <li><a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">Human mitochondrial genetics</a></li> <li><a href="/wiki/Mitochondrial_disease" title="Mitochondrial disease">Mitochondrial disease</a></li> <li><a href="/wiki/Mitochondrial_DNA_(journal)" title="Mitochondrial DNA (journal)">Mitochondrial DNA (journal)</a></li> <li><a href="/wiki/Mitochondrial_Eve" title="Mitochondrial Eve">Mitochondrial Eve</a></li> <li><a href="/wiki/Cambridge_Reference_Sequence" title="Cambridge Reference Sequence">Mitochondrial rCRS</a></li> <li><a href="/wiki/Paternal_mtDNA_transmission" title="Paternal mtDNA transmission">Paternal mtDNA transmission</a></li> <li><a href="/wiki/Single_origin_theory" class="mw-redirect" title="Single origin theory">Single origin theory</a></li> <li><a href="/wiki/Supercluster_(genetic)" title="Supercluster (genetic)">Supercluster (genetic)</a></li> <li><a href="/wiki/TIM/TOM_complex" title="TIM/TOM complex">TIM/TOM complex</a></li></ul></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Mitochondrial_DNA&action=edit&section=31" title="Edit section: References">edit</a>]</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"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Siekevitz-1"><a href="#cite_ref-Siekevitz_1-0">^</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 a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFSiekevitz1957" 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"Powerhouse of the cell". <a href="/wiki/Scientific_American" title="Scientific American">Scientific American</a>. 197 (1): 131–40. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1957SciAm.197a.131S">1957SciAm.197a.131S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fscientificamerican0757-131">10.1038/scientificamerican0757-131</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scientific+American&rft.atitle=Powerhouse+of+the+cell&rft.volume=197&rft.issue=1&rft.pages=131-40&rft.date=1957&rft_id=info%3Adoi%2F10.1038%2Fscientificamerican0757-131&rft_id=info%3Abibcode%2F1957SciAm.197a.131S&rft.aulast=Siekevitz&rft.aufirst=P&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrial+DNA" class="Z3988"> </li> <li id="cite_note-pmid15157274-2"><a href="#cite_ref-pmid15157274_2-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFIborraKimuraCook2004" class="citation journal cs1">Iborra FJ, Kimura H, Cook PR (May 2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC425603">"The functional organization of mitochondrial genomes in human cells"</a>. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Genetics&rft.atitle=An+atlas+of+mitochondrial+DNA+genotype-phenotype+associations+in+the+UK+Biobank&rft.volume=53&rft.issue=7&rft.pages=982-993&rft.date=2021-07&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7611844%23id-name%3DPMC&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A234768578%23id-name%3DS2CID&rft_id=info%3Apmid%2F34002094&rft_id=info%3Adoi%2F10.1038%2Fs41588-021-00868-1&rft.aulast=Yonova-Doing&rft.aufirst=E&rft.au=Calabrese%2C+C&rft.au=Gomez-Duran%2C+A&rft.au=Schon%2C+K&rft.au=Wei%2C+W&rft.au=Karthikeyan%2C+S&rft.au=Chinnery%2C+PF&rft.au=Howson%2C+JM&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7611844&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMitochondrial+DNA" class="Z3988"> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a 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.navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Chromosomes" title="Template:Chromosomes"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Chromosomes" title="Template talk:Chromosomes"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Chromosomes" title="Special:EditPage/Template:Chromosomes"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_genome" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_genome" title="Human genome">Human genome</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nuclear_DNA" title="Nuclear DNA">Nuclear genome</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%"><a href="/wiki/Autosome" title="Autosome">Autosome</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/Chromosome_1" title="Chromosome 1">1</a></li> <li><a href="/wiki/Chromosome_2" title="Chromosome 2">2</a></li> <li><a href="/wiki/Chromosome_3" title="Chromosome 3">3</a></li> <li><a href="/wiki/Chromosome_4" title="Chromosome 4">4</a></li> <li><a href="/wiki/Chromosome_5" title="Chromosome 5">5</a></li> <li><a href="/wiki/Chromosome_6" title="Chromosome 6">6</a></li> <li><a href="/wiki/Chromosome_7" title="Chromosome 7">7</a></li> <li><a href="/wiki/Chromosome_8" title="Chromosome 8">8</a></li> <li><a href="/wiki/Chromosome_9" title="Chromosome 9">9</a></li> <li><a href="/wiki/Chromosome_10" title="Chromosome 10">10</a></li> <li><a href="/wiki/Chromosome_11" title="Chromosome 11">11</a></li> <li><a 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style="padding:0 0.25em"> <ul><li><a href="/wiki/X_chromosome" title="X chromosome">X</a></li> <li><a href="/wiki/Y_chromosome" title="Y chromosome">Y</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Mitochondrial genome</a></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/Human_mitochondrial_genetics" title="Human mitochondrial genetics">Human mitochondrial DNA</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</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/Human_genome" title="Human genome">Human genome</a> <ul><li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li></ul></li> <li><a href="/wiki/List_of_human_genes" class="mw-redirect" title="List of human genes">List of human genes</a></li> <li><a href="/wiki/Archaeogenetics#Human_archaeology" title="Archaeogenetics">Human archaeology</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"><style data-mw-deduplicate="TemplateStyles:r886047488">.mw-parser-output .nobold{font-weight:normal}</style></div><div role="navigation" class="navbox" aria-labelledby="Types_of_nucleic_acids" 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" 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:Nucleic_acids" title="Template:Nucleic acids"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nucleic_acids" title="Template talk:Nucleic acids"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nucleic_acids" title="Special:EditPage/Template:Nucleic acids"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Types_of_nucleic_acids" style="font-size:114%;margin:0 4em">Types of <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids </a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">Constituents</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/Nucleobase" class="mw-redirect" title="Nucleobase">Nucleobases</a></li> <li><a href="/wiki/Nucleoside" title="Nucleoside">Nucleosides</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotides</a></li> <li><a href="/wiki/Deoxyribonucleotide" title="Deoxyribonucleotide">Deoxynucleotides</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/RNA" title="RNA">Ribonucleic acids </a> (coding, <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding</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%;text-align: center;"><a href="/wiki/RNA#In_translation" title="RNA">Translational</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/Messenger_RNA" title="Messenger RNA">Messenger </a> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">precursor, heterogenous nuclear</a></li></ul></li> <li><a href="/wiki/Nucleoside-modified_messenger_RNA" title="Nucleoside-modified messenger RNA">modified Messenger</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer </a></li> <li><a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">Ribosomal </a></li> <li><a href="/wiki/Transfer-messenger_RNA" title="Transfer-messenger RNA">Transfer-messenger </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/RNA#Regulatory_RNAs" title="RNA">Regulatory</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/RNA_interference" title="RNA interference">Interferential </a> <ul><li><a href="/wiki/MicroRNA" title="MicroRNA">Micro </a></li> <li><a href="/wiki/Small_interfering_RNA" title="Small interfering RNA">Small interfering </a></li> <li><a href="/wiki/Piwi-interacting_RNA" title="Piwi-interacting RNA">Piwi-interacting</a></li></ul></li> <li><a href="/wiki/Antisense_RNA" title="Antisense RNA">Antisense </a></li> <li><a href="/wiki/RNA#In_RNA_processing" title="RNA">Processual</a> <ul><li><a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">Small nuclear </a></li> <li><a href="/wiki/Small_nucleolar_RNA" title="Small nucleolar RNA">Small nucleolar </a></li> <li><a href="/wiki/Small_Cajal_body-specific_RNA" title="Small Cajal body-specific RNA">Small Cajal Body RNAs</a></li> <li><a href="/wiki/Y_RNA" title="Y RNA">Y RNA</a></li></ul></li> <li><a href="/wiki/Enhancer_RNA" title="Enhancer RNA">Enhancer RNAs </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Others</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/Guide_RNA" title="Guide RNA">Guide </a></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Small_hairpin_RNA" class="mw-redirect" title="Small hairpin RNA">Small hairpin </a></li> <li><a href="/wiki/Small_temporal_RNA" title="Small temporal RNA">Small temporal </a></li> <li><a href="/wiki/Trans-acting_siRNA" title="Trans-acting siRNA">Trans-acting small interfering</a></li> <li><a href="/wiki/Subgenomic_mRNA" title="Subgenomic mRNA">Subgenomic messenger</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/DNA" title="DNA">Deoxyribonucleic acids </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/Organellar_DNA" title="Organellar DNA">Organellar</a> <ul><li><a href="/wiki/Chloroplast_DNA" title="Chloroplast DNA">Chloroplast </a></li> <li><a class="mw-selflink selflink">Mitochondrial</a></li></ul></li> <li><a href="/wiki/Complementary_DNA" title="Complementary DNA">Complementary </a></li> <li><a href="/wiki/Deoxyribozyme" title="Deoxyribozyme">Deoxyribozyme</a></li> <li><a href="/wiki/Genomic_DNA" title="Genomic DNA">Genomic </a></li> <li><a href="/wiki/Hachimoji_DNA" title="Hachimoji DNA">Hachimoji</a></li> <li><a href="/wiki/Multicopy_single-stranded_DNA" title="Multicopy single-stranded DNA">Multicopy single-stranded </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid_analogue" title="Nucleic acid analogue">Analogues</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/Xeno_nucleic_acid" title="Xeno nucleic acid">Xeno </a> <ul><li><a href="/wiki/Glycol_nucleic_acid" title="Glycol nucleic acid">Glycol </a></li> <li><a href="/wiki/Threose_nucleic_acid" title="Threose nucleic acid">Threose </a></li> <li><a href="/wiki/Hexose" title="Hexose">Hexose</a></li></ul></li> <li><a href="/wiki/Locked_nucleic_acid" title="Locked nucleic acid">Locked </a></li> <li><a href="/wiki/Peptide_nucleic_acid" title="Peptide nucleic acid">Peptide </a></li> <li><a href="/wiki/Morpholino" title="Morpholino">Morpholino</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Cloning_vector" title="Cloning vector">Cloning vectors</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/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Lambda_phage" title="Lambda phage">Lambda phage</a></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Artificial chromosomes</a> <ul><li><a href="/wiki/P1-derived_artificial_chromosome" title="P1-derived artificial chromosome">P1-derived </a></li> <li><a href="/wiki/Bacterial_artificial_chromosome" title="Bacterial artificial chromosome">Bacterial </a></li> <li><a href="/wiki/Yeast_artificial_chromosome" title="Yeast artificial chromosome">Yeast </a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Human</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align: center;"><div> <ul><li><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /> <a href="/wiki/Category:Nucleic_acids" title="Category:Nucleic acids">Category</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="Mitochondrial_proteins" 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"><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:Mitochondrial_proteins" title="Template:Mitochondrial proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Mitochondrial_proteins" title="Template talk:Mitochondrial proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Mitochondrial_proteins" title="Special:EditPage/Template:Mitochondrial proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Mitochondrial_proteins" style="font-size:114%;margin:0 4em"><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrial</a> proteins</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Outer_mitochondrial_membrane" class="mw-redirect" title="Outer mitochondrial membrane">Outer 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/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 class="mw-selflink selflink">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"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a>: National <a href="https://www.wikidata.org/wiki/Q27075#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></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 rel="nofollow" class="external text" href="https://d-nb.info/gnd/4202301-4">Germany</a></li><li><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh92006022">United States</a></li><li><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007539431305171">Israel</a></li></ul></div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;justify-content:center;align-items:baseline}.mw-parser-output .portal-bar-bordered{padding:0 2em;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em auto 0}.mw-parser-output .portal-bar-related{font-size:100%;justify-content:flex-start}.mw-parser-output .portal-bar-unbordered{padding:0 1.7em;margin-left:0}.mw-parser-output .portal-bar-header{margin:0 1em 0 0.5em;flex:0 0 auto;min-height:24px}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;padding:0.15em 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