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class="vector-toc-list"> </ul> </li> <li id="toc-RNA_methylation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_methylation"> <div class="vector-toc-text"> 2.2 RNA methylation </div> </a> <ul id="toc-RNA_methylation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Histone_modifications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Histone_modifications"> <div class="vector-toc-text"> 2.3 Histone modifications </div> </a> <ul id="toc-Histone_modifications-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_transcripts" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_transcripts"> <div class="vector-toc-text"> 2.4 RNA transcripts </div> </a> <ul id="toc-RNA_transcripts-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-MicroRNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#MicroRNAs"> <div class="vector-toc-text"> 2.5 MicroRNAs </div> </a> <ul id="toc-MicroRNAs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-mRNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#mRNA"> <div class="vector-toc-text"> 2.6 mRNA </div> </a> <ul id="toc-mRNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-sRNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#sRNAs"> <div class="vector-toc-text"> 2.7 sRNAs </div> </a> <ul id="toc-sRNAs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Long_non-coding_RNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Long_non-coding_RNAs"> <div class="vector-toc-text"> 2.8 Long non-coding RNAs </div> </a> <ul id="toc-Long_non-coding_RNAs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Prions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Prions"> <div class="vector-toc-text"> 2.9 Prions </div> </a> <ul id="toc-Prions-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Molecular_basis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Molecular_basis"> <div class="vector-toc-text"> 3 Molecular basis </div> </a> <button aria-controls="toc-Molecular_basis-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Molecular basis subsection </button> <ul id="toc-Molecular_basis-sublist" class="vector-toc-list"> <li id="toc-DNA_damage" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_damage"> <div class="vector-toc-text"> 3.1 DNA damage </div> </a> <ul id="toc-DNA_damage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_repair" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_repair"> <div class="vector-toc-text"> 3.2 DNA repair </div> </a> <ul id="toc-DNA_repair-sublist" class="vector-toc-list"> <li id="toc-Repair_of_oxidative_DNA_damage_can_alter_epigenetic_markers" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Repair_of_oxidative_DNA_damage_can_alter_epigenetic_markers"> <div class="vector-toc-text"> 3.2.1 Repair of oxidative DNA damage can alter epigenetic markers </div> </a> <ul id="toc-Repair_of_oxidative_DNA_damage_can_alter_epigenetic_markers-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Homologous_recombinational_repair_alters_epigenetic_markers" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Homologous_recombinational_repair_alters_epigenetic_markers"> <div class="vector-toc-text"> 3.2.2 Homologous recombinational repair alters epigenetic markers </div> </a> <ul id="toc-Homologous_recombinational_repair_alters_epigenetic_markers-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-homologous_end_joining_can_cause_some_epigenetic_marker_alterations" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Non-homologous_end_joining_can_cause_some_epigenetic_marker_alterations"> <div class="vector-toc-text"> 3.2.3 Non-homologous end joining can cause some epigenetic marker alterations </div> </a> <ul id="toc-Non-homologous_end_joining_can_cause_some_epigenetic_marker_alterations-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Techniques_used_to_study_epigenetics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Techniques_used_to_study_epigenetics"> <div class="vector-toc-text"> 3.3 Techniques used to study epigenetics </div> </a> <ul id="toc-Techniques_used_to_study_epigenetics-sublist" class="vector-toc-list"> <li id="toc-Chromatin_Immunoprecipitation" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Chromatin_Immunoprecipitation"> <div class="vector-toc-text"> 3.3.1 Chromatin Immunoprecipitation </div> </a> <ul id="toc-Chromatin_Immunoprecipitation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Fluorescent_in_situ_hybridization" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Fluorescent_in_situ_hybridization"> <div class="vector-toc-text"> 3.3.2 Fluorescent in situ hybridization </div> </a> <ul id="toc-Fluorescent_in_situ_hybridization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Methylation-sensitive_restriction_enzymes" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Methylation-sensitive_restriction_enzymes"> <div class="vector-toc-text"> 3.3.3 Methylation-sensitive restriction enzymes </div> </a> <ul id="toc-Methylation-sensitive_restriction_enzymes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bisulfite_sequencing" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Bisulfite_sequencing"> <div class="vector-toc-text"> 3.3.4 Bisulfite sequencing </div> </a> <ul id="toc-Bisulfite_sequencing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nanopore_sequencing" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Nanopore_sequencing"> <div class="vector-toc-text"> 3.3.5 Nanopore sequencing </div> </a> <ul id="toc-Nanopore_sequencing-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Structural_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structural_inheritance"> <div class="vector-toc-text"> 3.4 Structural inheritance </div> </a> <ul id="toc-Structural_inheritance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nucleosome_positioning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nucleosome_positioning"> <div class="vector-toc-text"> 3.5 Nucleosome positioning </div> </a> <ul id="toc-Nucleosome_positioning-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Histone_variants" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Histone_variants"> <div class="vector-toc-text"> 3.6 Histone variants </div> </a> <ul id="toc-Histone_variants-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genomic_architecture" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genomic_architecture"> <div class="vector-toc-text"> 3.7 Genomic architecture </div> </a> <ul id="toc-Genomic_architecture-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Functions_and_consequences" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Functions_and_consequences"> <div class="vector-toc-text"> 4 Functions and consequences </div> </a> <button aria-controls="toc-Functions_and_consequences-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Functions and consequences subsection </button> <ul id="toc-Functions_and_consequences-sublist" class="vector-toc-list"> <li id="toc-In_the_brain" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_the_brain"> <div class="vector-toc-text"> 4.1 In the brain </div> </a> <ul id="toc-In_the_brain-sublist" class="vector-toc-list"> <li id="toc-Memory" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Memory"> <div class="vector-toc-text"> 4.1.1 Memory </div> </a> <ul id="toc-Memory-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Aging" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Aging"> <div class="vector-toc-text"> 4.1.2 Aging </div> </a> <ul id="toc-Aging-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_and_general" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Other_and_general"> <div class="vector-toc-text"> 4.1.3 Other and general </div> </a> <ul id="toc-Other_and_general-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Development"> <div class="vector-toc-text"> 4.2 Development </div> </a> <ul id="toc-Development-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transgenerational" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transgenerational"> <div class="vector-toc-text"> 4.3 Transgenerational </div> </a> <ul id="toc-Transgenerational-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Epigenetics_in_bacteria" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Epigenetics_in_bacteria"> <div class="vector-toc-text"> 5 Epigenetics in bacteria </div> </a> <ul id="toc-Epigenetics_in_bacteria-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Medicine" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Medicine"> <div class="vector-toc-text"> 6 Medicine </div> </a> <button aria-controls="toc-Medicine-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Medicine subsection </button> <ul id="toc-Medicine-sublist" class="vector-toc-list"> <li id="toc-Twins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Twins"> <div class="vector-toc-text"> 6.1 Twins </div> </a> <ul id="toc-Twins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genomic_imprinting" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genomic_imprinting"> <div class="vector-toc-text"> 6.2 Genomic imprinting </div> </a> <ul id="toc-Genomic_imprinting-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Examples_of_drugs_altering_gene_expression_from_epigenetic_events" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Examples_of_drugs_altering_gene_expression_from_epigenetic_events"> <div class="vector-toc-text"> 6.3 Examples of drugs altering gene expression from epigenetic events </div> </a> <ul id="toc-Examples_of_drugs_altering_gene_expression_from_epigenetic_events-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Addiction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Addiction"> <div class="vector-toc-text"> 6.4 Addiction </div> </a> <ul id="toc-Addiction-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Research" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Research"> <div class="vector-toc-text"> 7 Research </div> </a> <button aria-controls="toc-Research-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Research subsection </button> <ul id="toc-Research-sublist" class="vector-toc-list"> <li id="toc-Epigenome_editing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Epigenome_editing"> <div class="vector-toc-text"> 7.1 Epigenome editing </div> </a> <ul id="toc-Epigenome_editing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-CpG_sites,_SNPs_and_biological_traits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#CpG_sites,_SNPs_and_biological_traits"> <div class="vector-toc-text"> 7.2 CpG sites, SNPs and biological traits </div> </a> <ul id="toc-CpG_sites,_SNPs_and_biological_traits-sublist" class="vector-toc-list"> <li id="toc-UBASH3B_locus" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#UBASH3B_locus"> <div class="vector-toc-text"> 7.2.1 UBASH3B locus </div> </a> <ul id="toc-UBASH3B_locus-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NFKBIE_locus" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#NFKBIE_locus"> <div class="vector-toc-text"> 7.2.2 NFKBIE locus </div> </a> <ul id="toc-NFKBIE_locus-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-FADS1_locus" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#FADS1_locus"> <div class="vector-toc-text"> 7.2.3 FADS1 locus </div> </a> <ul id="toc-FADS1_locus-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Pseudoscience" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Pseudoscience"> <div class="vector-toc-text"> 8 Pseudoscience </div> </a> <ul id="toc-Pseudoscience-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 9 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"> 10 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> 11 Further reading </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 12 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">Epigenetics</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-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B9%D9%84%D9%85_%D8%A7%D9%84%D8%AA%D8%AE%D9%84%D9%82" title="علم التخلق – Arabic" lang="ar" hreflang="ar" data-title="علم التخلق" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Epigenetika" title="Epigenetika – Azerbaijani" lang="az" hreflang="az" data-title="Epigenetika" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%85%E0%A6%A7%E0%A6%BF%E0%A6%AC%E0%A6%82%E0%A6%B6%E0%A6%BE%E0%A6%A3%E0%A7%81%E0%A6%AC%E0%A6%BF%E0%A6%9C%E0%A7%8D%E0%A6%9E%E0%A6%BE%E0%A6%A8" title="অধিবংশাণুবিজ্ঞান – Bangla" lang="bn" hreflang="bn" data-title="অধিবংশাণুবিজ্ঞান" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%95%D0%BF%D0%B8%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Епигенетика – Bulgarian" lang="bg" hreflang="bg" data-title="Епигенетика" 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/Epigenetika" title="Epigenetika – Bosnian" lang="bs" hreflang="bs" data-title="Epigenetika" 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/Epigen%C3%A8tica" title="Epigenètica – Catalan" lang="ca" hreflang="ca" data-title="Epigenètica" 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/Epigenetika" title="Epigenetika – Czech" lang="cs" hreflang="cs" data-title="Epigenetika" 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/Epigenetik" title="Epigenetik – Danish" lang="da" hreflang="da" data-title="Epigenetik" 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/Epigenetik" title="Epigenetik – German" lang="de" hreflang="de" data-title="Epigenetik" 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/Epigeneetika" title="Epigeneetika – Estonian" lang="et" hreflang="et" data-title="Epigeneetika" 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%95%CF%80%CE%B9%CE%B3%CE%B5%CE%BD%CE%B5%CF%84%CE%B9%CE%BA%CE%AE" title="Επιγενετική – Greek" lang="el" hreflang="el" data-title="Επιγενετική" 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/Epigen%C3%A9tica" title="Epigenética – Spanish" lang="es" hreflang="es" data-title="Epigenética" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Epigenetiko" title="Epigenetiko – Esperanto" lang="eo" hreflang="eo" data-title="Epigenetiko" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Epigenetika" title="Epigenetika – Basque" lang="eu" hreflang="eu" data-title="Epigenetika" 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%A7%D9%BE%DB%8C%E2%80%8C%DA%98%D9%86%D8%AA%DB%8C%DA%A9" title="اپی‌ژنتیک – Persian" lang="fa" hreflang="fa" data-title="اپی‌ژنتیک" 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/%C3%89pig%C3%A9n%C3%A9tique" title="Épigénétique – French" lang="fr" hreflang="fr" data-title="Épigénétique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Eipig%C3%A9ineola%C3%ADocht" title="Eipigéineolaíocht – Irish" lang="ga" hreflang="ga" data-title="Eipigéineolaíocht" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Epixen%C3%A9tica" title="Epixenética – Galician" lang="gl" hreflang="gl" data-title="Epixenética" 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/%ED%9B%84%EC%84%B1%EC%9C%A0%EC%A0%84%ED%95%99" title="후성유전학 – Korean" lang="ko" hreflang="ko" data-title="후성유전학" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Epigenetika" title="Epigenetika – Croatian" lang="hr" hreflang="hr" data-title="Epigenetika" 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/Epigenetika" title="Epigenetika – Indonesian" lang="id" hreflang="id" data-title="Epigenetika" 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/Epigenetica" title="Epigenetica – Italian" lang="it" hreflang="it" data-title="Epigenetica" 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%90%D7%A4%D7%99%D7%92%D7%A0%D7%98%D7%99%D7%A7%D7%94" title="אפיגנטיקה – Hebrew" lang="he" hreflang="he" data-title="אפיגנטיקה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%8E%E0%B2%AA%E0%B2%BF%E0%B2%9C%E0%B3%86%E0%B2%A8%E0%B3%86%E0%B2%9F%E0%B2%BF%E0%B2%95%E0%B3%8D%E0%B2%B8%E0%B3%8D%E2%80%8C" title="ಎಪಿಜೆನೆಟಿಕ್ಸ್‌ – Kannada" lang="kn" hreflang="kn" data-title="ಎಪಿಜೆನೆಟಿಕ್ಸ್‌" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Epi%C4%A3en%C4%93tika" title="Epiģenētika – Latvian" lang="lv" hreflang="lv" data-title="Epiģenētika" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Epigenetika" title="Epigenetika – Hungarian" lang="hu" hreflang="hu" data-title="Epigenetika" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%95%D0%BF%D0%B8%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Епигенетика – Macedonian" lang="mk" hreflang="mk" data-title="Епигенетика" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%8E%E0%B4%AA%E0%B4%BF%E0%B4%9C%E0%B5%86%E0%B4%A8%E0%B5%86%E0%B4%B1%E0%B5%8D%E0%B4%B1%E0%B4%BF%E0%B4%95%E0%B5%8D%E0%B4%B8%E0%B5%8D" title="എപിജെനെറ്റിക്സ് – Malayalam" lang="ml" hreflang="ml" data-title="എപിജെനെറ്റിക്സ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%86%E0%A4%A8%E0%A5%81%E0%A4%B5%E0%A4%BE%E0%A4%82%E0%A4%B6%E0%A4%BF%E0%A4%95_%E0%A4%9C%E0%A4%A8%E0%A5%81%E0%A4%95%E0%A4%B6%E0%A4%BE%E0%A4%B8%E0%A5%8D%E0%A4%A4%E0%A5%8D%E0%A4%B0" title="आनुवांशिक जनुकशास्त्र – Marathi" lang="mr" hreflang="mr" data-title="आनुवांशिक जनुकशास्त्र" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target">मराठी</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Epigenetik" title="Epigenetik – Malay" lang="ms" hreflang="ms" data-title="Epigenetik" 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%AD%D0%BF%D0%B8%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA" title="Эпигенетик – Mongolian" lang="mn" hreflang="mn" data-title="Эпигенетик" 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/Epigenetica" title="Epigenetica – Dutch" lang="nl" hreflang="nl" data-title="Epigenetica" 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%82%A8%E3%83%94%E3%82%B8%E3%82%A7%E3%83%8D%E3%83%86%E3%82%A3%E3%82%AF%E3%82%B9" title="エピジェネティクス – Japanese" lang="ja" hreflang="ja" data-title="エピジェネティクス" 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-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Epigenetikk" title="Epigenetikk – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Epigenetikk" 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/%D8%A7%D9%BE%D9%8A_%DA%98%D9%86%D9%BC%DB%8C%DA%A9" title="اپي ژنټیک – Pashto" lang="ps" hreflang="ps" data-title="اپي ژنټیک" 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/Epigenetyka" title="Epigenetyka – Polish" lang="pl" hreflang="pl" data-title="Epigenetyka" 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/Epigen%C3%A9tica" title="Epigenética – Portuguese" lang="pt" hreflang="pt" data-title="Epigenética" 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/Epigenetic%C4%83" title="Epigenetică – Romanian" lang="ro" hreflang="ro" data-title="Epigenetică" 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%AD%D0%BF%D0%B8%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Эпигенетика – Russian" lang="ru" hreflang="ru" data-title="Эпигенетика" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Epigjenetika" title="Epigjenetika – Albanian" lang="sq" hreflang="sq" data-title="Epigjenetika" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target">Shqip</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Epigenetics" title="Epigenetics – Simple English" lang="en-simple" hreflang="en-simple" data-title="Epigenetics" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Epigenetika" title="Epigenetika – Slovenian" lang="sl" hreflang="sl" data-title="Epigenetika" 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%95%D0%BF%D0%B8%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Епигенетика – Serbian" lang="sr" hreflang="sr" data-title="Епигенетика" 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/Epigenetika" title="Epigenetika – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Epigenetika" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Epigenetiikka" title="Epigenetiikka – Finnish" lang="fi" hreflang="fi" data-title="Epigenetiikka" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Epigenetik" title="Epigenetik – Swedish" lang="sv" hreflang="sv" data-title="Epigenetik" 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%AE%E0%AF%87%E0%AE%B2%E0%AF%8D%E0%AE%AE%E0%AE%B0%E0%AE%AA%E0%AE%BF%E0%AE%AF%E0%AE%B2%E0%AF%8D" title="மேல்மரபியல் – Tamil" lang="ta" hreflang="ta" data-title="மேல்மரபியல்" 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%AD%E0%B8%B5%E0%B8%9E%E0%B8%B5%E0%B9%80%E0%B8%88%E0%B9%80%E0%B8%99%E0%B8%95%E0%B8%B4%E0%B8%81%E0%B8%AA%E0%B9%8C" title="อีพีเจเนติกส์ – Thai" lang="th" hreflang="th" data-title="อีพีเจเนติกส์" 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/Epigenetik" title="Epigenetik – Turkish" lang="tr" hreflang="tr" data-title="Epigenetik" 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%95%D0%BF%D1%96%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Епігенетика – Ukrainian" lang="uk" hreflang="uk" data-title="Епігенетика" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%A8%D8%A7%D9%84%D8%A7%D9%88%D8%B1%D8%A7%D8%AB%DB%8C%D8%A7%D8%AA" title="بالاوراثیات – Urdu" lang="ur" hreflang="ur" data-title="بالاوراثیات" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Di_truy%E1%BB%81n_h%E1%BB%8Dc_bi%E1%BB%83u_sinh" title="Di truyền học biểu sinh – Vietnamese" lang="vi" hreflang="vi" data-title="Di truyền học biểu sinh" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E8%A1%A8%E8%A7%80%E9%81%BA%E5%82%B3%E5%AD%B8" title="表觀遺傳學 – Cantonese" lang="yue" hreflang="yue" data-title="表觀遺傳學" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target">粵語</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%A1%A8%E8%A7%80%E9%81%BA%E5%82%B3%E5%AD%B8" title="表觀遺傳學 – Chinese" lang="zh" hreflang="zh" data-title="表觀遺傳學" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li> </ul> <div 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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 other uses, see <a href="/wiki/Epigenetic_(disambiguation)" class="mw-disambig" title="Epigenetic (disambiguation)">Epigenetic (disambiguation)</a>.</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Epigenetic_mechanisms.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Epigenetic_mechanisms.png/449px-Epigenetic_mechanisms.png" decoding="async" width="449" height="305" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Epigenetic_mechanisms.png/674px-Epigenetic_mechanisms.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Epigenetic_mechanisms.png/898px-Epigenetic_mechanisms.png 2x" data-file-width="2187" data-file-height="1488" /></a><figcaption>Epigenetic mechanisms</figcaption></figure> In <a href="/wiki/Biology" title="Biology">biology</a>, epigenetics is the study of <a href="/wiki/Heritability" title="Heritability">heritable traits</a>, or a stable change of cell function, that happen without changes to the <a href="/wiki/DNA_sequence" class="mw-redirect" title="DNA sequence">DNA sequence</a>.<a href="#cite_note-Epigenetics_2009_review-1">[1]</a> The <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> prefix <a href="https://en.wiktionary.org/wiki/epi-" class="extiw" title="wikt:epi-">epi-</a> (<a href="https://en.wiktionary.org/wiki/%E1%BC%90%CF%80%CE%B9-#Ancient_Greek" class="extiw" title="wikt:ἐπι-">ἐπι-</a> "over, outside of, around") in epigenetics implies features that are "on top of" or "in addition to" the traditional (DNA sequence based) <a href="/wiki/Gene" title="Gene">genetic</a> mechanism of inheritance.<a href="#cite_note-science-2">[2]</a> Epigenetics usually involves a change that is not erased by cell division, and affects the <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulation of gene expression</a>.<a href="#cite_note-3">[3]</a> Such effects on <a href="/wiki/Cell_(biology)" title="Cell (biology)">cellular</a> and <a href="/wiki/Physiology" title="Physiology">physiological</a> <a href="/wiki/Phenotypic_trait" title="Phenotypic trait">phenotypic traits</a> may result from <a href="/wiki/Environment_(biophysical)" class="mw-redirect" title="Environment (biophysical)">environmental</a> factors, or be part of normal development. Epigenetic factors can also lead to cancer.<a href="#cite_note-4">[4]</a> The term also refers to the mechanism of changes: functionally relevant alterations to the <a href="/wiki/Genome" title="Genome">genome</a> that do not involve mutation of the <a href="/wiki/Nucleotide_sequence" class="mw-redirect" title="Nucleotide sequence">nucleotide sequence</a>. Examples of mechanisms that produce such changes are <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> and <a href="/wiki/Histone_modification" class="mw-redirect" title="Histone modification">histone modification</a>, each of which alters how genes are expressed without altering the underlying <a href="/wiki/DNA" title="DNA">DNA</a> sequence.<a href="#cite_note-5">[5]</a> Further, non-coding RNA sequences have been shown to play a key role in the regulation of gene expression.<a href="#cite_note-6">[6]</a> Gene expression can be controlled through the action of <a href="/wiki/Repressor_protein" class="mw-redirect" title="Repressor protein">repressor proteins</a> that attach to <a href="/wiki/Silencer_(DNA)" class="mw-redirect" title="Silencer (DNA)">silencer</a> regions of the DNA. These epigenetic changes may last through <a href="/wiki/Cell_division" title="Cell division">cell divisions</a> for the duration of the cell's life, and may also last for multiple generations, even though they do not involve changes in the underlying DNA sequence of the organism;<a href="#cite_note-pmid17522671-7">[7]</a> instead, non-genetic factors cause the organism's genes to behave (or "express themselves") differently.<a href="#cite_note-8">[8]</a> One example of an epigenetic change in <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> biology is the process of <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">cellular differentiation</a>. During <a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenesis</a>, <a href="/wiki/Totipotent" class="mw-redirect" title="Totipotent">totipotent</a> <a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">stem cells</a> become the various <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a> <a href="/wiki/Cell_line" class="mw-redirect" title="Cell line">cell lines</a> of the <a href="/wiki/Embryo" title="Embryo">embryo</a>, which in turn become fully differentiated cells. In other words, as a single fertilized <a href="/wiki/Egg_cell" title="Egg cell">egg cell</a> – the <a href="/wiki/Zygote" title="Zygote">zygote</a> – continues to <a href="/wiki/Mitosis" title="Mitosis">divide</a>, the resulting daughter cells change into all the different cell types in an organism, including <a href="/wiki/Neurons" class="mw-redirect" title="Neurons">neurons</a>, <a href="/wiki/Muscle_cells" class="mw-redirect" title="Muscle cells">muscle cells</a>, <a href="/wiki/Epithelium" title="Epithelium">epithelium</a>, <a href="/wiki/Endothelium" title="Endothelium">endothelium</a> of <a href="/wiki/Blood_vessels" class="mw-redirect" title="Blood vessels">blood vessels</a>, etc., by activating some genes while inhibiting the expression of others.<a href="#cite_note-pmid17522676-9">[9]</a> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Definitions">Definitions</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=1" title="Edit section: Definitions">edit</a>]</div> The term epigenesis has a generic meaning of "extra growth" that has been used in English since the 17th century.<a href="#cite_note-10">[10]</a> In scientific publications, the term epigenetics started to appear in the 1930s (see Fig. on the right). However, its contemporary meaning emerged only in the 1990s.<a href="#cite_note-Moore_2015-11">[11]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:EpigenByYear_1.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/EpigenByYear_1.png/220px-EpigenByYear_1.png" decoding="async" width="220" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/EpigenByYear_1.png/330px-EpigenByYear_1.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/64/EpigenByYear_1.png/440px-EpigenByYear_1.png 2x" data-file-width="3705" data-file-height="2272" /></a><figcaption>Number of patent families and non-patent documents with the term "epigenetic*" by publication year</figcaption></figure> A definition of the concept of epigenetic trait as a "stably heritable phenotype resulting from changes in a chromosome without alterations in the DNA sequence" was formulated at a <a href="/wiki/Cold_Spring_Harbor_Laboratory" title="Cold Spring Harbor Laboratory">Cold Spring Harbor</a> meeting in 2008,<a href="#cite_note-pmid19339683-12">[12]</a> although alternate definitions that include non-heritable traits are still being used widely.<a href="#cite_note-NIH-13">[13]</a> <div class="mw-heading mw-heading3"><h3 id="Waddington's_canalisation,_1940s">Waddington's canalisation, 1940s</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=2" title="Edit section: Waddington's canalisation, 1940s">edit</a>]</div> The hypothesis of epigenetic changes affecting the expression of <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> was put forth by the Russian biologist <a href="/wiki/Nikolai_Koltsov" title="Nikolai Koltsov">Nikolai Koltsov</a>.<a href="#cite_note-14">[14]</a> From the generic meaning, and the associated adjective epigenetic, British embryologist <a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a> coined the term epigenetics in 1942 as pertaining to <a href="/wiki/Epigenesis_(biology)" title="Epigenesis (biology)">epigenesis</a>, in parallel to <a href="/wiki/Valentin_Haecker" title="Valentin Haecker">Valentin Haecker</a>'s 'phenogenetics' (Phänogenetik).<a href="#cite_note-waddington-15">[15]</a> Epigenesis in the context of the biology of that period referred to the <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a> of cells from their initial <a href="/wiki/Totipotent" class="mw-redirect" title="Totipotent">totipotent</a> state during <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a>.<a href="#cite_note-16">[16]</a> When Waddington coined the term, the physical nature of <a href="/wiki/Gene" title="Gene">genes</a> and their role in heredity was not known. He used it instead as a conceptual model of how genetic components might interact with their surroundings to produce a <a href="/wiki/Phenotype" title="Phenotype">phenotype</a>; he used the phrase "<a href="/wiki/Epigenetic_landscape" class="mw-redirect" title="Epigenetic landscape">epigenetic landscape</a>" as a metaphor for <a href="/wiki/Morphogenesis" title="Morphogenesis">biological development</a>. Waddington held that cell fates were established during development in a process he called <a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">canalisation</a> much as a marble rolls down to the point of <a href="/wiki/Local_optimum" class="mw-redirect" title="Local optimum">lowest local elevation</a>.<a href="#cite_note-Waddington2014-17">[17]</a> Waddington suggested visualising increasing irreversibility of cell type differentiation as ridges rising between the valleys where the marbles (analogous to cells) are travelling.<a href="#cite_note-18">[18]</a> In recent times, Waddington's notion of the epigenetic landscape has been rigorously formalized in the context of the <a href="/wiki/System_dynamics" title="System dynamics">systems dynamics</a> state approach to the study of cell-fate.<a href="#cite_note-19">[19]</a><a href="#cite_note-sciencedirect.com-20">[20]</a> Cell-fate determination is predicted to exhibit certain dynamics, such as attractor-convergence (the attractor can be an equilibrium point, limit cycle or <a href="/wiki/Strange_attractor" class="mw-redirect" title="Strange attractor">strange attractor</a>) or oscillatory.<a href="#cite_note-sciencedirect.com-20">[20]</a> <div class="mw-heading mw-heading3"><h3 id="Contemporary">Contemporary</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=3" title="Edit section: Contemporary">edit</a>]</div> <a href="/wiki/Robin_Holliday" title="Robin Holliday">Robin Holliday</a> defined in 1990 epigenetics as "the study of the mechanisms of temporal and spatial control of gene activity during the development of complex organisms."<a href="#cite_note-pmid2265224-21">[21]</a> More recent usage of the word in biology follows stricter definitions. As defined by <a href="/wiki/Arthur_Riggs_(geneticist)" title="Arthur Riggs (geneticist)">Arthur Riggs</a> and colleagues, it is "the study of <a href="/wiki/Mitosis" title="Mitosis">mitotically</a> and/or <a href="/wiki/Meiosis" title="Meiosis">meiotically</a> heritable changes in gene function that cannot be explained by changes in DNA sequence."<a href="#cite_note-isbn0-87969-490-4-22">[22]</a> The term has also been used, however, to describe processes which have not been demonstrated to be heritable, such as some forms of histone modification. Consequently, there are attempts to redefine "epigenetics" in broader terms that would avoid the constraints of requiring <a href="/wiki/Heritability" title="Heritability">heritability</a>. For example, <a href="/wiki/Adrian_Peter_Bird" class="mw-redirect" title="Adrian Peter Bird">Adrian Bird</a> defined epigenetics as "the structural adaptation of chromosomal regions so as to register, signal or perpetuate altered activity states."<a href="#cite_note-pmid17522671-7">[7]</a> This definition would be inclusive of transient modifications associated with <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> or <a href="/wiki/Cell-cycle" class="mw-redirect" title="Cell-cycle">cell-cycle</a> phases as well as stable changes maintained across multiple cell generations, but exclude others such as templating of membrane architecture and <a href="/wiki/Prions" class="mw-redirect" title="Prions">prions</a> unless they impinge on chromosome function. Such redefinitions however are not universally accepted and are still subject to debate.<a href="#cite_note-nature2008-23">[23]</a> The <a href="/wiki/National_Institutes_of_Health" title="National Institutes of Health">NIH</a> "Roadmap Epigenomics Project", which ran from 2008 to 2017, uses the following definition: "For purposes of this program, epigenetics refers to both heritable changes in gene activity and <a href="/wiki/Gene_expression" title="Gene expression">expression</a> (in the progeny of cells or of individuals) and also stable, long-term alterations in the transcriptional potential of a cell that are not necessarily heritable."<a href="#cite_note-24">[24]</a> In 2008, a consensus definition of the epigenetic trait, a "stably heritable phenotype resulting from changes in a chromosome without alterations in the DNA sequence," was made at a <a href="/wiki/Cold_Spring_Harbor_Laboratory" title="Cold Spring Harbor Laboratory">Cold Spring Harbor</a> meeting.<a href="#cite_note-pmid19339683-12">[12]</a> The similarity of the word to "genetics" has generated many parallel usages. The "<a href="/wiki/Epigenome" title="Epigenome">epigenome</a>" is a parallel to the word "<a href="/wiki/Genome" title="Genome">genome</a>", referring to the overall epigenetic state of a cell, and <a href="/wiki/Epigenomics" title="Epigenomics">epigenomics</a> refers to global analyses of epigenetic changes across the entire genome.<a href="#cite_note-NIH-13">[13]</a> The phrase "<a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>" has also been adapted – the "<a href="/wiki/Epigenetic_code" title="Epigenetic code">epigenetic code</a>" has been used to describe the set of epigenetic features that create different phenotypes in different cells from the same underlying DNA sequence. Taken to its extreme, the "epigenetic code" could represent the total state of the cell, with the position of each molecule accounted for in an epigenomic map, a diagrammatic representation of the gene expression, DNA methylation and histone modification status of a particular genomic region. More typically, the term is used in reference to systematic efforts to measure specific, relevant forms of epigenetic information such as the <a href="/wiki/Histone_code_hypothesis" class="mw-redirect" title="Histone code hypothesis">histone code</a> or <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> patterns.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading2"><h2 id="Mechanisms">Mechanisms</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=4" title="Edit section: Mechanisms">edit</a>]</div> <a href="/wiki/Covalent" class="mw-redirect" title="Covalent">Covalent</a> modification of either DNA (e.g. cytosine methylation and hydroxymethylation) or of histone proteins (e.g. lysine acetylation, lysine and arginine methylation, serine and threonine phosphorylation, and lysine ubiquitination and sumoylation) play central roles in many types of epigenetic inheritance. Therefore, the word "epigenetics" is sometimes used as a synonym for these processes. However, this can be misleading. Chromatin remodeling is not always inherited, and not all epigenetic inheritance involves chromatin remodeling.<a href="#cite_note-pmid17407749-25">[25]</a> In 2019, a further lysine modification appeared in the scientific literature linking epigenetics modification to cell metabolism, i.e. lactylation<a href="#cite_note-26">[26]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nucleosome_1KX5_2.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Nucleosome_1KX5_2.png/220px-Nucleosome_1KX5_2.png" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Nucleosome_1KX5_2.png/330px-Nucleosome_1KX5_2.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Nucleosome_1KX5_2.png/440px-Nucleosome_1KX5_2.png 2x" data-file-width="1024" data-file-height="1024" /></a><figcaption>DNA associates with histone proteins to form chromatin.</figcaption></figure> Because the <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> of a cell or individual is affected by which of its genes are transcribed, heritable <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription states</a> can give rise to epigenetic effects. There are several layers of regulation of <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>. One way that genes are regulated is through the remodeling of chromatin. Chromatin is the complex of DNA and the <a href="/wiki/Histone" title="Histone">histone</a> proteins with which it associates. If the way that DNA is wrapped around the histones changes, gene expression can change as well. Chromatin remodeling is accomplished through two main mechanisms: <ol><li>The first way is <a href="/wiki/Post_translational_modification" class="mw-redirect" title="Post translational modification">post translational modification</a> of the amino acids that make up histone proteins. Histone proteins are made up of long chains of amino acids. If the amino acids that are in the chain are changed, the shape of the histone might be modified. DNA is not completely unwound during replication. It is possible, then, that the modified histones may be carried into each new copy of the DNA. Once there, these histones may act as templates, initiating the surrounding new histones to be shaped in the new manner. By altering the shape of the histones around them, these modified histones would ensure that a lineage-specific transcription program is maintained after cell division.</li> <li>The second way is the addition of methyl groups to the DNA, mostly at <a href="/wiki/CpG_site" title="CpG site">CpG sites</a>, to convert <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> to <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a>. 5-Methylcytosine performs much like a regular cytosine, pairing with a guanine in double-stranded DNA. However, when methylated cytosines are present in <a href="/wiki/CpG_site" title="CpG site">CpG sites</a> in the <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions of genes, the genes are often repressed.<a href="#cite_note-pmid30619465-27">[27]</a><a href="#cite_note-pmid31399642-28">[28]</a> When methylated cytosines are present in <a href="/wiki/CpG_site" title="CpG site">CpG sites</a> in the gene body (in the <a href="/wiki/Coding_region" title="Coding region">coding region</a> excluding the transcription start site) expression of the gene is often enhanced. Transcription of a gene usually depends on a <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factor</a> binding to a (10 base or less) <a href="/wiki/Recognition_sequence" title="Recognition sequence">recognition sequence</a> at the enhancer that interacts with the promoter region of that gene (<a href="/wiki/Gene_expression#Enhancers,_transcription_factors,_mediator_complex_and_DNA_loops_in_mammalian_transcription" title="Gene expression">Gene expression#Enhancers, transcription factors, mediator complex and DNA loops in mammalian transcription</a>).<a href="#cite_note-pmid22868264-29">[29]</a> About 22% of transcription factors are inhibited from binding when the recognition sequence has a methylated cytosine. In addition, presence of methylated cytosines at a promoter region can attract <a href="/wiki/Methyl-CpG-binding_domain" title="Methyl-CpG-binding domain">methyl-CpG-binding domain</a> (MBD) proteins. All MBDs interact with <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a> remodeling and <a href="/wiki/Histone_deacetylase" title="Histone deacetylase">histone deacetylase</a> complexes, which leads to gene silencing. In addition, another covalent modification involving methylated cytosine is its <a href="/wiki/DNA_demethylation" title="DNA demethylation">demethylation</a> by <a href="/wiki/TET_enzymes" title="TET enzymes">TET enzymes</a>. Hundreds of such demethylations occur, for instance, during <a href="/wiki/DNA_demethylation#Learnking_and_memory" title="DNA demethylation">learning and memory</a> forming events in <a href="/wiki/Neuron" title="Neuron">neurons</a>.<a href="#cite_note-pmid28620075-30">[30]</a><a href="#cite_note-Bernstein-31">[31]</a></li></ol> There is frequently a reciprocal relationship between DNA methylation and histone lysine methylation.<a href="#cite_note-Rose-32">[32]</a> For instance, the <a href="/wiki/Methyl-CpG-binding_domain" title="Methyl-CpG-binding domain">methyl binding domain protein MBD1</a>, attracted to and associating with <a href="/wiki/5-Methylcytosine" title="5-Methylcytosine">methylated cytosine</a> in a DNA <a href="/wiki/CpG_site" title="CpG site">CpG site</a>, can also associate with H3K9 <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">methyltransferase</a> activity to methylate histone 3 at lysine 9. On the other hand, DNA maintenance methylation by <a href="/wiki/DNMT1" title="DNMT1">DNMT1</a> appears to partly rely on recognition of histone methylation on the nucleosome present at the DNA site to carry out cytosine methylation on newly synthesized DNA.<a href="#cite_note-Rose-32">[32]</a> There is further crosstalk between DNA methylation carried out by <a href="/wiki/DNMT3A" class="mw-redirect" title="DNMT3A">DNMT3A</a> and <a href="/wiki/DNMT3B" title="DNMT3B">DNMT3B</a> and histone methylation so that there is a correlation between the genome-wide distribution of DNA methylation and histone methylation.<a href="#cite_note-Li2021-33">[33]</a> Mechanisms of heritability of histone state are not well understood; however, much is known about the mechanism of heritability of DNA methylation state during cell division and differentiation. Heritability of methylation state depends on certain enzymes (such as <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNMT1</a>) that have a higher affinity for 5-methylcytosine than for cytosine. If this enzyme reaches a "hemimethylated" portion of DNA (where 5-methylcytosine is in only one of the two DNA strands) the enzyme will methylate the other half. However, it is now known that DNMT1 physically interacts with the protein <a href="/wiki/UHRF1" title="UHRF1">UHRF1</a>. UHRF1 has been recently recognized as essential for DNMT1-mediated maintenance of DNA methylation. UHRF1 is the protein that specifically recognizes hemi-methylated DNA, therefore bringing DNMT1 to its substrate to maintain DNA methylation.<a href="#cite_note-Li2021-33">[33]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Histone_tails_set_for_transcriptional_activation.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Histone_tails_set_for_transcriptional_activation.jpg/220px-Histone_tails_set_for_transcriptional_activation.jpg" decoding="async" width="220" height="195" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Histone_tails_set_for_transcriptional_activation.jpg/330px-Histone_tails_set_for_transcriptional_activation.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Histone_tails_set_for_transcriptional_activation.jpg/440px-Histone_tails_set_for_transcriptional_activation.jpg 2x" data-file-width="2346" data-file-height="2082" /></a><figcaption>Some acetylations and some methylations of lysines (symbol K) are activation signals for transcription when present on a <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a>, as shown in the top figure. Some methylations on lysines or arginine (R) are repression signals for transcription when present on a <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a>, as shown in the bottom figure. <a href="/wiki/Nucleosome" title="Nucleosome">Nucleosomes</a> consist of four pairs of <a href="/wiki/Histone" title="Histone">histone</a> proteins in a tightly assembled core region plus up to 30% of each histone remaining in a loosely organized tail<a href="#cite_note-pmid33133421-34">[34]</a> (only one tail of each pair is shown). DNA is wrapped around the histone core proteins in <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>. The lysines (K) are designated with a number showing their position as, for instance (K4), indicating lysine as the 4th amino acid from the amino (N) end of the tail in the histone protein. <a href="/wiki/Methylation" title="Methylation">Methylations</a> [Me], and <a href="/wiki/Acetylation" title="Acetylation">acetylations</a> [Ac] are common <a href="/wiki/Post-translational_modification" title="Post-translational modification">post-translational modifications</a> on the lysines of the histone tails.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Histone_tails_set_for_transcriptional_repression.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Histone_tails_set_for_transcriptional_repression.jpg/220px-Histone_tails_set_for_transcriptional_repression.jpg" decoding="async" width="220" height="195" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Histone_tails_set_for_transcriptional_repression.jpg/330px-Histone_tails_set_for_transcriptional_repression.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/42/Histone_tails_set_for_transcriptional_repression.jpg/440px-Histone_tails_set_for_transcriptional_repression.jpg 2x" data-file-width="2315" data-file-height="2048" /></a><figcaption></figcaption></figure> Although histone modifications occur throughout the entire sequence, the unstructured N-termini of histones (called histone tails) are particularly highly modified. These modifications include <a href="/wiki/Acetylation" title="Acetylation">acetylation</a>, <a href="/wiki/Methylation" title="Methylation">methylation</a>, <a href="/wiki/Ubiquitylation" class="mw-redirect" title="Ubiquitylation">ubiquitylation</a>, <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a>, <a href="/wiki/Sumoylation" class="mw-redirect" title="Sumoylation">sumoylation</a>, ribosylation and citrullination. Acetylation is the most highly studied of these modifications. For example, acetylation of the K14 and K9 <a href="/wiki/Lysine" title="Lysine">lysines</a> of the tail of histone H3 by histone acetyltransferase enzymes (HATs) is generally related to transcriptional competence<a href="#cite_note-35">[35]</a> (see Figure). One mode of thinking is that this tendency of acetylation to be associated with "active" transcription is biophysical in nature. Because it normally has a positively charged nitrogen at its end, lysine can bind the negatively charged phosphates of the DNA backbone. The acetylation event converts the positively charged amine group on the side chain into a neutral amide linkage. This removes the positive charge, thus loosening the DNA from the histone. When this occurs, complexes like <a href="/wiki/SWI/SNF" title="SWI/SNF">SWI/SNF</a> and other transcriptional factors can bind to the DNA and allow transcription to occur. This is the "cis" model of the epigenetic function. In other words, changes to the histone tails have a direct effect on the DNA itself.<a href="#cite_note-36">[36]</a> Another model of epigenetic function is the "trans" model. In this model, changes to the histone tails act indirectly on the DNA. For example, lysine acetylation may create a binding site for chromatin-modifying enzymes (or transcription machinery as well). This chromatin remodeler can then cause changes to the state of the chromatin. Indeed, a bromodomain – a protein domain that specifically binds acetyl-lysine – is found in many enzymes that help activate transcription, including the <a href="/wiki/SWI/SNF" title="SWI/SNF">SWI/SNF</a> complex. It may be that acetylation acts in this and the previous way to aid in transcriptional activation. The idea that modifications act as docking modules for related factors is borne out by <a href="/wiki/Histone_methylation" title="Histone methylation">histone methylation</a> as well. Methylation of lysine 9 of histone H3 has long been associated with constitutively transcriptionally silent chromatin (constitutive <a href="/wiki/Heterochromatin" title="Heterochromatin">heterochromatin</a>) (see bottom Figure). It has been determined that a chromodomain (a domain that specifically binds methyl-lysine) in the transcriptionally repressive protein <a href="/wiki/Heterochromatin_Protein_1" class="mw-redirect" title="Heterochromatin Protein 1">HP1</a> recruits HP1 to K9 methylated regions. One example that seems to refute this biophysical model for methylation is that tri-methylation of histone H3 at lysine 4 is strongly associated with (and required for full) transcriptional activation (see top Figure). Tri-methylation, in this case, would introduce a fixed positive charge on the tail. It has been shown that the histone lysine methyltransferase (KMT) is responsible for this methylation activity in the pattern of histones H3 & H4. This enzyme utilizes a catalytically active site called the SET domain (Suppressor of variegation, Enhancer of Zeste, Trithorax). The SET domain is a 130-amino acid sequence involved in modulating gene activities. This domain has been demonstrated to bind to the histone tail and causes the methylation of the histone.<a href="#cite_note-pmid9487389-37">[37]</a> Differing histone modifications are likely to function in differing ways; acetylation at one position is likely to function differently from acetylation at another position. Also, multiple modifications may occur at the same time, and these modifications may work together to change the behavior of the <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a>. The idea that multiple dynamic modifications regulate gene transcription in a systematic and reproducible way is called the <a href="/wiki/Histone_code" title="Histone code">histone code</a>, although the idea that histone state can be read linearly as a digital information carrier has been largely debunked. One of the best-understood systems that orchestrate chromatin-based silencing is the <a href="/wiki/SIR_protein" class="mw-redirect" title="SIR protein">SIR protein</a> based silencing of the yeast hidden mating-type loci HML and HMR. <div class="mw-heading mw-heading3"><h3 id="DNA_methylation">DNA methylation</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=5" title="Edit section: DNA methylation">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/Methylation" title="Methylation">Methylation</a></div> <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> frequently occurs in repeated sequences, and helps to suppress the expression and mobility of '<a href="/wiki/Transposable_elements" class="mw-redirect" title="Transposable elements">transposable elements</a>':<a href="#cite_note-slotkin2007-38">[38]</a> Because <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a> can be spontaneously deaminated (replacing nitrogen by oxygen) to <a href="/wiki/Thymidine" title="Thymidine">thymidine</a>, CpG sites are frequently mutated and become rare in the genome, except at <a href="/wiki/CpG_islands" class="mw-redirect" title="CpG islands">CpG islands</a> where they remain unmethylated. Epigenetic changes of this type thus have the potential to direct increased frequencies of permanent genetic mutation. DNA methylation patterns are known to be established and modified in response to environmental factors by a complex interplay of at least three independent <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferases</a>, DNMT1, DNMT3A, and DNMT3B, the loss of any of which is lethal in mice.<a href="#cite_note-li92-39">[39]</a> DNMT1 is the most abundant methyltransferase in somatic cells,<a href="#cite_note-robertson99-40">[40]</a> localizes to replication foci,<a href="#cite_note-leonhardt92-41">[41]</a> has a 10–40-fold preference for hemimethylated DNA and interacts with the <a href="/wiki/Proliferating_cell_nuclear_antigen" title="Proliferating cell nuclear antigen">proliferating cell nuclear antigen</a> (PCNA).<a href="#cite_note-chuang97-42">[42]</a> By preferentially modifying hemimethylated DNA, DNMT1 transfers patterns of methylation to a newly synthesized strand after <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>, and therefore is often referred to as the 'maintenance' methyltransferase.<a href="#cite_note-robertson00-43">[43]</a> DNMT1 is essential for proper embryonic development, imprinting and X-inactivation.<a href="#cite_note-li92-39">[39]</a><a href="#cite_note-li93-44">[44]</a> To emphasize the difference of this molecular mechanism of inheritance from the canonical Watson-Crick base-pairing mechanism of transmission of genetic information, the term 'Epigenetic templating' was introduced.<a href="#cite_note-45">[45]</a> Furthermore, in addition to the maintenance and transmission of methylated DNA states, the same principle could work in the maintenance and transmission of histone modifications and even cytoplasmic (<a href="/wiki/Structural_inheritance" title="Structural inheritance">structural</a>) heritable states.<a href="#cite_note-pmid18419815-46">[46]</a> <div class="mw-heading mw-heading3"><h3 id="RNA_methylation">RNA methylation</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=6" title="Edit section: RNA methylation">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/Methylation" title="Methylation">Methylation</a></div> RNA methylation of N6-methyladenosine (m6A) as the most abundant eukaryotic RNA modification has recently been recognized as an important gene regulatory mechanism.<a href="#cite_note-pmid32300195-47">[47]</a> <div class="mw-heading mw-heading3"><h3 id="Histone_modifications">Histone modifications</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=7" title="Edit section: Histone modifications">edit</a>]</div> Histones H3 and H4 can also be manipulated through demethylation using <a href="/wiki/Histone_lysine_demethylase" class="mw-redirect" title="Histone lysine demethylase">histone lysine demethylase</a> (KDM). This recently identified enzyme has a catalytically active site called the Jumonji domain (JmjC). The demethylation occurs when JmjC utilizes multiple cofactors to hydroxylate the methyl group, thereby removing it. JmjC is capable of demethylating mono-, di-, and tri-methylated substrates.<a href="#cite_note-pmid19234061-48">[48]</a> Chromosomal regions can adopt stable and heritable alternative states resulting in bistable gene expression without changes to the DNA sequence. Epigenetic control is often associated with alternative <a href="/wiki/Covalent_modification" class="mw-redirect" title="Covalent modification">covalent modifications</a> of histones.<a href="#cite_note-Rosenfeld_2009-49">[49]</a> The stability and heritability of states of larger chromosomal regions are suggested to involve positive feedback where modified <a href="/wiki/Nucleosome" title="Nucleosome">nucleosomes</a> recruit enzymes that similarly modify nearby nucleosomes.<a href="#cite_note-50">[50]</a> A simplified stochastic model for this type of epigenetics is found here.<a href="#cite_note-51">[51]</a><a href="#cite_note-pmid17512413-52">[52]</a> It has been suggested that chromatin-based transcriptional regulation could be mediated by the effect of small RNAs. <a href="/wiki/Small_interfering_RNA" title="Small interfering RNA">Small interfering RNAs</a> can modulate transcriptional gene expression via epigenetic modulation of targeted <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoters</a>.<a href="#cite_note-Morris-53">[53]</a> <div class="mw-heading mw-heading3"><h3 id="RNA_transcripts">RNA transcripts</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=8" title="Edit section: RNA transcripts">edit</a>]</div> Sometimes a gene, after being turned on, transcribes a product that (directly or indirectly) maintains the activity of that gene. For example, <a href="/wiki/Hnf4" class="mw-redirect" title="Hnf4">Hnf4</a> and <a href="/wiki/MyoD" title="MyoD">MyoD</a> enhance the transcription of many liver-specific and muscle-specific genes, respectively, including their own, through the <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factor</a> activity of the <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a> they encode. RNA signalling includes differential recruitment of a hierarchy of generic chromatin modifying complexes and DNA methyltransferases to specific loci by RNAs during differentiation and development.<a href="#cite_note-pmid19154003-54">[54]</a> Other epigenetic changes are mediated by the production of <a href="/wiki/Alternative_splicing" title="Alternative splicing">different splice forms</a> of <a href="/wiki/RNA" title="RNA">RNA</a>, or by formation of double-stranded RNA (<a href="/wiki/RNAi" class="mw-redirect" title="RNAi">RNAi</a>). Descendants of the cell in which the gene was turned on will inherit this activity, even if the original stimulus for gene-activation is no longer present. These genes are often turned on or off by <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a>, although in some systems where <a href="/wiki/Syncytia" class="mw-redirect" title="Syncytia">syncytia</a> or <a href="/wiki/Gap_junction" title="Gap junction">gap junctions</a> are important, RNA may spread directly to other cells or nuclei by <a href="/wiki/Diffusion" title="Diffusion">diffusion</a>. A large amount of RNA and protein is contributed to the <a href="/wiki/Zygote" title="Zygote">zygote</a> by the mother during <a href="/wiki/Oogenesis" title="Oogenesis">oogenesis</a> or via <a href="/wiki/Nurse_cell" title="Nurse cell">nurse cells</a>, resulting in <a href="/wiki/Maternal_effect" title="Maternal effect">maternal effect</a> phenotypes. A smaller quantity of sperm RNA is transmitted from the father, but there is recent evidence that this epigenetic information can lead to visible changes in several generations of offspring.<a href="#cite_note-choi06-55">[55]</a> <div class="mw-heading mw-heading3"><h3 id="MicroRNAs">MicroRNAs</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=9" title="Edit section: MicroRNAs">edit</a>]</div> <a href="/wiki/MicroRNA" title="MicroRNA">MicroRNAs</a> (miRNAs) are members of <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding RNAs</a> that range in size from 17 to 25 nucleotides. miRNAs regulate a large variety of biological functions in plants and animals.<a href="#cite_note-Wang-56">[56]</a> So far, in 2013, about 2000 miRNAs have been discovered in humans and these can be found online in a miRNA database.<a href="#cite_note-57">[57]</a> Each miRNA expressed in a cell may target about 100 to 200 messenger RNAs(mRNAs) that it downregulates.<a href="#cite_note-58">[58]</a> Most of the downregulation of mRNAs occurs by causing the decay of the targeted mRNA, while some downregulation occurs at the level of translation into protein.<a href="#cite_note-59">[59]</a> It appears that about 60% of human protein coding genes are regulated by miRNAs.<a href="#cite_note-60">[60]</a> Many miRNAs are epigenetically regulated. About 50% of miRNA genes are associated with <a href="/wiki/CpG_island" class="mw-redirect" title="CpG island">CpG islands</a>,<a href="#cite_note-Wang-56">[56]</a> that may be repressed by epigenetic methylation. Transcription from methylated CpG islands is strongly and heritably repressed.<a href="#cite_note-61">[61]</a> Other miRNAs are epigenetically regulated by either histone modifications or by combined DNA methylation and histone modification.<a href="#cite_note-Wang-56">[56]</a> <div class="mw-heading mw-heading3"><h3 id="mRNA">mRNA</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=10" title="Edit section: mRNA">edit</a>]</div> In 2011, it was demonstrated that the <a href="/wiki/Methylation" title="Methylation">methylation</a> of <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a> plays a critical role in human <a href="/wiki/Energy_balance_(biology)" class="mw-redirect" title="Energy balance (biology)">energy homeostasis</a>. The obesity-associated <a href="/wiki/FTO_gene" title="FTO gene">FTO gene</a> is shown to be able to <a href="/wiki/Demethylate" class="mw-redirect" title="Demethylate">demethylate</a> <a href="/wiki/N6-methyladenosine" class="mw-redirect" title="N6-methyladenosine">N6-methyladenosine</a> in RNA.<a href="#cite_note-62">[62]</a><a href="#cite_note-63">[63]</a> <div class="mw-heading mw-heading3"><h3 id="sRNAs">sRNAs</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=11" title="Edit section: sRNAs">edit</a>]</div> <a href="/wiki/Bacterial_small_RNA" title="Bacterial small RNA">sRNAs</a> are small (50–250 nucleotides), highly structured, non-coding RNA fragments found in bacteria. They control gene expression including <a href="/wiki/Virulence" title="Virulence">virulence</a> genes in pathogens and are viewed as new targets in the fight against drug-resistant bacteria.<a href="#cite_note-64">[64]</a> They play an important role in many biological processes, binding to mRNA and protein targets in prokaryotes. Their phylogenetic analyses, for example through sRNA–mRNA target interactions or protein <a href="/wiki/Hfq_binding_sRNA" title="Hfq binding sRNA">binding properties</a>, are used to build comprehensive databases.<a href="#cite_note-65">[65]</a> sRNA-<a href="/wiki/Gene_map" class="mw-redirect" title="Gene map">gene maps</a> based on their targets in microbial genomes are also constructed.<a href="#cite_note-66">[66]</a> <div class="mw-heading mw-heading3"><h3 id="Long_non-coding_RNAs">Long non-coding RNAs</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=12" title="Edit section: Long non-coding RNAs">edit</a>]</div> Numerous investigations have demonstrated the pivotal involvement of long non-coding RNAs (lncRNAs) in the regulation of gene expression and chromosomal modifications, thereby exerting significant control over cellular differentiation. These long non-coding RNAs also contribute to genomic imprinting and the inactivation of the X chromosome.<a href="#cite_note-67">[67]</a> In invertebrates such as social insects of honey bees, long non-coding RNAs are detected as a possible epigenetic mechanism via allele-specific genes underlying aggression via reciprocal crosses.<a href="#cite_note-68">[68]</a> <div class="mw-heading mw-heading3"><h3 id="Prions">Prions</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=13" title="Edit section: Prions">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/Fungal_prions" class="mw-redirect" title="Fungal prions">Fungal prions</a></div> <a href="/wiki/Prion" title="Prion">Prions</a> are <a href="/wiki/Infection" title="Infection">infectious</a> forms of <a href="/wiki/Protein" title="Protein">proteins</a>. In general, proteins fold into discrete units that perform distinct cellular functions, but some proteins are also capable of forming an infectious conformational state known as a prion. Although often viewed in the context of <a href="/wiki/Transmissible_spongiform_encephalopathy" title="Transmissible spongiform encephalopathy">infectious disease</a>, prions are more loosely defined by their ability to catalytically convert other native state versions of the same protein to an infectious conformational state. It is in this latter sense that they can be viewed as epigenetic agents capable of inducing a phenotypic change without a modification of the genome.<a href="#cite_note-69">[69]</a> <a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prions</a> are considered by some to be epigenetic because the infectious phenotype caused by the prion can be inherited without modification of the genome. <a href="/wiki/PSI_(prion)" class="mw-redirect" title="PSI (prion)">PSI+</a> and URE3, discovered in <a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">yeast</a> in 1965 and 1971, are the two best studied of this type of prion.<a href="#cite_note-70">[70]</a><a href="#cite_note-pmid5573734-71">[71]</a> Prions can have a phenotypic effect through the sequestration of protein in aggregates, thereby reducing that protein's activity. In PSI+ cells, the loss of the Sup35 protein (which is involved in termination of translation) causes ribosomes to have a higher rate of read-through of stop <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>, an effect that results in suppression of <a href="/wiki/Nonsense_mutation" title="Nonsense mutation">nonsense mutations</a> in other genes.<a href="#cite_note-pmid225301-72">[72]</a> The ability of Sup35 to form prions may be a conserved trait. It could confer an adaptive advantage by giving cells the ability to <a href="/wiki/Evolutionary_capacitance" title="Evolutionary capacitance">switch into a PSI+ state</a> and express dormant genetic features normally terminated by stop codon mutations.<a href="#cite_note-pmid11028992-73">[73]</a><a href="#cite_note-pmid15931169-74">[74]</a><a href="#cite_note-75">[75]</a><a href="#cite_note-76">[76]</a> Prion-based epigenetics has also been observed in <a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a>.<a href="#cite_note-77">[77]</a> <div class="mw-heading mw-heading2"><h2 id="Molecular_basis">Molecular basis</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=14" title="Edit section: Molecular basis">edit</a>]</div> Epigenetic changes modify the activation of certain genes, but not the genetic code sequence of DNA.<a href="#cite_note-Topart-78">[78]</a> The microstructure (not code) of DNA itself or the associated <a href="/wiki/Chromatin" title="Chromatin">chromatin</a> proteins may be modified, causing activation or silencing. This mechanism enables differentiated cells in a multicellular organism to express only the genes that are necessary for their own activity. Epigenetic changes are preserved when cells divide. Most epigenetic changes only occur within the course of one individual organism's lifetime; however, these epigenetic changes can be transmitted to the organism's offspring through a process called <a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">transgenerational epigenetic inheritance</a>. Moreover, if gene inactivation occurs in a sperm or egg cell that results in fertilization, this epigenetic modification may also be transferred to the next generation.<a href="#cite_note-pmid17320501-79">[79]</a> Specific epigenetic processes include <a href="/wiki/Paramutation" title="Paramutation">paramutation</a>, <a href="/wiki/Bookmarking" title="Bookmarking">bookmarking</a>, <a href="/wiki/Imprinting_(genetics)" class="mw-redirect" title="Imprinting (genetics)">imprinting</a>, <a href="/wiki/Gene_silencing" title="Gene silencing">gene silencing</a>, <a href="/wiki/X-inactivation" title="X-inactivation">X chromosome inactivation</a>, <a href="/wiki/Position_effect" title="Position effect">position effect</a>, <a href="/wiki/DNA_methylation_reprogramming" class="mw-redirect" title="DNA methylation reprogramming">DNA methylation reprogramming</a>, <a href="/wiki/Transvection_(genetics)" title="Transvection (genetics)">transvection</a>, <a href="/wiki/Maternal_effect" title="Maternal effect">maternal effects</a>, the progress of <a href="/wiki/Carcinogenesis" title="Carcinogenesis">carcinogenesis</a>, many effects of <a href="/wiki/Teratogen" class="mw-redirect" title="Teratogen">teratogens</a>, regulation of <a href="/wiki/Histone" title="Histone">histone</a> modifications and <a href="/wiki/Heterochromatin" title="Heterochromatin">heterochromatin</a>, and technical limitations affecting <a href="/wiki/Parthenogenesis" title="Parthenogenesis">parthenogenesis</a> and <a href="/wiki/Cloning" title="Cloning">cloning</a>.<a href="#cite_note-80">[80]</a><a href="#cite_note-81">[81]</a><a href="#cite_note-82">[82]</a> <div class="mw-heading mw-heading3"><h3 id="DNA_damage">DNA damage</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=15" title="Edit section: DNA damage">edit</a>]</div> DNA damage can also cause epigenetic changes.<a href="#cite_note-83">[83]</a><a href="#cite_note-84">[84]</a><a href="#cite_note-85">[85]</a> DNA damage is very frequent, occurring on average about 60,000 times a day per cell of the human body (see <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage (naturally occurring)</a>). These damages are largely repaired, however, epigenetic changes can still remain at the site of DNA repair.<a href="#cite_note-86">[86]</a> In particular, a double strand break in DNA can initiate unprogrammed epigenetic gene silencing both by causing DNA methylation as well as by promoting silencing types of histone modifications (chromatin remodeling - see next section).<a href="#cite_note-87">[87]</a> In addition, the enzyme <a href="/wiki/Poly_ADP_ribose_polymerase" class="mw-redirect" title="Poly ADP ribose polymerase">Parp1 (poly(ADP)-ribose polymerase)</a> and its product poly(ADP)-ribose (PAR) accumulate at sites of DNA damage as part of the repair process.<a href="#cite_note-88">[88]</a> This accumulation, in turn, directs recruitment and activation of the chromatin remodeling protein, ALC1, that can cause <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a> remodeling.<a href="#cite_note-89">[89]</a> Nucleosome remodeling has been found to cause, for instance, epigenetic silencing of DNA repair gene MLH1.<a href="#cite_note-isbn0-87969-490-4-22">[22]</a><a href="#cite_note-90">[90]</a> DNA damaging chemicals, such as <a href="/wiki/Benzene" title="Benzene">benzene</a>, <a href="/wiki/Hydroquinone" title="Hydroquinone">hydroquinone</a>, <a href="/wiki/Styrene" title="Styrene">styrene</a>, <a href="/wiki/Carbon_tetrachloride" title="Carbon tetrachloride">carbon tetrachloride</a> and <a href="/wiki/Trichloroethylene" title="Trichloroethylene">trichloroethylene</a>, cause considerable hypomethylation of DNA, some through the activation of oxidative stress pathways.<a href="#cite_note-91">[91]</a> Foods are known to alter the epigenetics of rats on different diets.<a href="#cite_note-92">[92]</a> Some food components epigenetically increase the levels of DNA repair enzymes such as <a href="/wiki/O-6-methylguanine-DNA_methyltransferase" class="mw-redirect" title="O-6-methylguanine-DNA methyltransferase">MGMT</a> and <a href="/wiki/MLH1" title="MLH1">MLH1</a><a href="#cite_note-93">[93]</a> and <a href="/wiki/P53" title="P53">p53</a>.<a href="#cite_note-94">[94]</a> Other food components can reduce DNA damage, such as soy <a href="/wiki/Isoflavones" class="mw-redirect" title="Isoflavones">isoflavones</a>. In one study, markers for oxidative stress, such as modified nucleotides that can result from DNA damage, were decreased by a 3-week diet supplemented with soy.<a href="#cite_note-95">[95]</a> A decrease in oxidative DNA damage was also observed 2 h after consumption of <a href="/wiki/Anthocyanin" title="Anthocyanin">anthocyanin</a>-rich <a href="/wiki/Bilberry" title="Bilberry">bilberry</a> (<a href="/wiki/Vaccinium_myrtillus" title="Vaccinium myrtillus">Vaccinium myrtillius</a> L.) <a href="/wiki/Pomace" title="Pomace">pomace</a> extract.<a href="#cite_note-96">[96]</a> <div class="mw-heading mw-heading3"><h3 id="DNA_repair">DNA repair</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=16" title="Edit section: DNA repair">edit</a>]</div> Damage to DNA is very common and is constantly being repaired. Epigenetic alterations can accompany DNA repair of oxidative damage or double-strand breaks. In human cells, oxidative DNA damage occurs about 10,000 times a day and DNA double-strand breaks occur about 10 to 50 times a cell cycle in somatic replicating cells (see <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage (naturally occurring)</a>). The selective advantage of DNA repair is to allow the cell to survive in the face of DNA damage. The selective advantage of epigenetic alterations that occur with DNA repair is not clear.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading4"><h4 id="Repair_of_oxidative_DNA_damage_can_alter_epigenetic_markers">Repair of oxidative DNA damage can alter epigenetic markers</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=17" title="Edit section: Repair of oxidative DNA damage can alter epigenetic markers">edit</a>]</div> In the steady state (with endogenous damages occurring and being repaired), there are about 2,400 oxidatively damaged guanines that form <a href="/wiki/8-oxo-2%27-deoxyguanosine" class="mw-redirect" title="8-oxo-2'-deoxyguanosine">8-oxo-2'-deoxyguanosine</a> (8-OHdG) in the average mammalian cell DNA.<a href="#cite_note-pmid21163908-97">[97]</a> 8-OHdG constitutes about 5% of the oxidative damages commonly present in DNA.<a href="#cite_note-Hamilton-98">[98]</a> The oxidized guanines do not occur randomly among all guanines in DNA. There is a sequence preference for the guanine at a <a href="/wiki/DNA_methylation" title="DNA methylation">methylated</a> <a href="/wiki/CpG_site" title="CpG site">CpG site</a> (a cytosine followed by guanine along its <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">5' → 3' direction</a> and where the cytosine is methylated (5-mCpG)).<a href="#cite_note-pmid24571128-99">[99]</a> A 5-mCpG site has the lowest ionization potential for guanine oxidation.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure typeof="mw:File/Thumb"><a href="/wiki/File:Initiation_of_DNA_demethylation_at_a_CpG_site.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Initiation_of_DNA_demethylation_at_a_CpG_site.svg/200px-Initiation_of_DNA_demethylation_at_a_CpG_site.svg.png" decoding="async" width="200" height="230" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Initiation_of_DNA_demethylation_at_a_CpG_site.svg/300px-Initiation_of_DNA_demethylation_at_a_CpG_site.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Initiation_of_DNA_demethylation_at_a_CpG_site.svg/400px-Initiation_of_DNA_demethylation_at_a_CpG_site.svg.png 2x" data-file-width="643" data-file-height="739" /></a><figcaption>Initiation of <a href="/wiki/DNA_demethylation" title="DNA demethylation">DNA demethylation</a> at a <a href="/wiki/CpG_site" title="CpG site">CpG site</a>. In adult somatic cells DNA methylation typically occurs in the context of CpG dinucleotides (<a href="/wiki/CpG_sites" class="mw-redirect" title="CpG sites">CpG sites</a>), forming <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a>-pG, or 5mCpG. Reactive oxygen species (ROS) may attack guanine at the dinucleotide site, forming <a href="/wiki/8-oxo-2%27-deoxyguanosine" class="mw-redirect" title="8-oxo-2'-deoxyguanosine">8-hydroxy-2'-deoxyguanosine</a> (8-OHdG), and resulting in a 5mCp-8-OHdG dinucleotide site. The <a href="/wiki/Base_excision_repair" title="Base excision repair">base excision repair</a> enzyme <a href="/wiki/Oxoguanine_glycosylase" title="Oxoguanine glycosylase">OGG1</a> targets 8-OHdG and binds to the lesion without immediate excision. OGG1, present at a 5mCp-8-OHdG site recruits <a href="/wiki/Tet_methylcytosine_dioxygenase_1" title="Tet methylcytosine dioxygenase 1">TET1</a> and TET1 oxidizes the 5mC adjacent to the 8-OHdG. This initiates demethylation of 5mC.<a href="#cite_note-Zhou-100">[100]</a></figcaption></figure> Oxidized guanine has mispairing potential and is mutagenic.<a href="#cite_note-pmid31993111-101">[101]</a> <a href="/wiki/Oxoguanine_glycosylase" title="Oxoguanine glycosylase">Oxoguanine glycosylase</a> (OGG1) is the primary enzyme responsible for the excision of the oxidized guanine during DNA repair. OGG1 finds and binds to an 8-OHdG within a few seconds.<a href="#cite_note-pmid33171795-102">[102]</a> However, OGG1 does not immediately excise 8-OHdG. In HeLa cells half maximum removal of 8-OHdG occurs in 30 minutes,<a href="#cite_note-pmid15365186-103">[103]</a> and in irradiated mice, the 8-OHdGs induced in the mouse liver are removed with a half-life of 11 minutes.<a href="#cite_note-Hamilton-98">[98]</a> When OGG1 is present at an oxidized guanine within a methylated <a href="/wiki/CpG_site" title="CpG site">CpG site</a> it recruits <a href="/wiki/TET_enzymes" title="TET enzymes">TET1</a> to the 8-OHdG lesion (see Figure). This allows TET1 to demethylate an adjacent methylated cytosine. Demethylation of cytosine is an epigenetic alteration.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] As an example, when human mammary epithelial cells were treated with H2O2 for six hours, 8-OHdG increased about 3.5-fold in DNA and this caused about 80% demethylation of the 5-methylcytosines in the genome.<a href="#cite_note-Zhou-100">[100]</a> Demethylation of CpGs in a gene promoter by <a href="/wiki/TET_enzymes" title="TET enzymes">TET enzyme</a> activity increases transcription of the gene into messenger RNA.<a href="#cite_note-pmid24108092-104">[104]</a> In cells treated with H2O2, one particular gene was examined, <a href="/wiki/Beta-secretase_1" title="Beta-secretase 1">BACE1</a>.<a href="#cite_note-Zhou-100">[100]</a> The methylation level of the BACE1 <a href="/wiki/CpG_site#CpG_islands" title="CpG site">CpG island</a> was reduced (an epigenetic alteration) and this allowed about 6.5 fold increase of expression of BACE1 messenger RNA.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] While six-hour incubation with H2O2 causes considerable demethylation of 5-mCpG sites, shorter times of H2O2 incubation appear to promote other epigenetic alterations. Treatment of cells with H2O2 for 30 minutes causes the mismatch repair protein heterodimer MSH2-MSH6 to recruit DNA methyltransferase 1 (<a href="/wiki/DNMT1" title="DNMT1">DNMT1</a>) to sites of some kinds of oxidative DNA damage.<a href="#cite_note-pmid26186941-105">[105]</a> This could cause increased methylation of cytosines (epigenetic alterations) at these locations. Jiang et al.<a href="#cite_note-Jiang-106">[106]</a> treated <a href="/wiki/HEK_293_cells" title="HEK 293 cells">HEK 293 cells</a> with agents causing oxidative DNA damage, (<a href="/wiki/Potassium_bromate" title="Potassium bromate">potassium bromate</a> (KBrO3) or <a href="/wiki/Potassium_chromate" title="Potassium chromate">potassium chromate</a> (K2CrO4)). <a href="/wiki/Base_excision_repair" title="Base excision repair">Base excision repair</a> (BER) of oxidative damage occurred with the DNA repair enzyme <a href="/wiki/DNA_polymerase" title="DNA polymerase">polymerase beta</a> localizing to oxidized guanines. Polymerase beta is the main human polymerase in short-patch BER of oxidative DNA damage. Jiang et al.<a href="#cite_note-Jiang-106">[106]</a> also found that polymerase beta recruited the <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferase</a> protein DNMT3b to BER repair sites. They then evaluated the methylation pattern at the single nucleotide level in a small region of DNA including the <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> region and the early transcription region of the <a href="/wiki/BRCA1" title="BRCA1">BRCA1</a> gene. Oxidative DNA damage from bromate modulated the DNA methylation pattern (caused epigenetic alterations) at CpG sites within the region of DNA studied. In untreated cells, CpGs located at −189, −134, −29, −19, +16, and +19 of the BRCA1 gene had methylated cytosines (where numbering is from the <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> transcription start site, and negative numbers indicate nucleotides in the upstream <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> region). Bromate treatment-induced oxidation resulted in the loss of cytosine methylation at −189, −134, +16 and +19 while also leading to the formation of new methylation at the CpGs located at −80, −55, −21 and +8 after DNA repair was allowed. <div class="mw-heading mw-heading4"><h4 id="Homologous_recombinational_repair_alters_epigenetic_markers">Homologous recombinational repair alters epigenetic markers</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=18" title="Edit section: Homologous recombinational repair alters epigenetic markers">edit</a>]</div> At least four articles report the recruitment of <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferase 1 (DNMT1)</a> to sites of DNA double-strand breaks.<a href="#cite_note-pmid15956212-107">[107]</a><a href="#cite_note-Cuozzo-108">[108]</a><a href="#cite_note-pmid18704159-109">[109]</a><a href="#cite_note-pmid20940144-110">[110]</a> During <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombinational repair (HR)</a> of the double-strand break, the involvement of DNMT1 causes the two repaired strands of DNA to have different levels of methylated cytosines. One strand becomes frequently methylated at about 21 <a href="/wiki/CpG_site" title="CpG site">CpG sites</a> downstream of the repaired double-strand break. The other DNA strand loses methylation at about six CpG sites that were previously methylated downstream of the double-strand break, as well as losing methylation at about five CpG sites that were previously methylated upstream of the double-strand break. When the chromosome is replicated, this gives rise to one daughter chromosome that is heavily methylated downstream of the previous break site and one that is unmethylated in the region both upstream and downstream of the previous break site. With respect to the gene that was broken by the double-strand break, half of the progeny cells express that gene at a high level and in the other half of the progeny cells expression of that gene is repressed. When clones of these cells were maintained for three years, the new methylation patterns were maintained over that time period.<a href="#cite_note-pmid27629060-111">[111]</a> In mice with a CRISPR-mediated homology-directed recombination insertion in their genome there were a large number of increased methylations of CpG sites within the double-strand break-associated insertion.<a href="#cite_note-pmid33267773-112">[112]</a> <div class="mw-heading mw-heading4"><h4 id="Non-homologous_end_joining_can_cause_some_epigenetic_marker_alterations">Non-homologous end joining can cause some epigenetic marker alterations</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=19" title="Edit section: Non-homologous end joining can cause some epigenetic marker alterations">edit</a>]</div> <a href="/wiki/Non-homologous_end_joining" title="Non-homologous end joining">Non-homologous end joining</a> (NHEJ) repair of a double-strand break can cause a small number of demethylations of pre-existing cytosine DNA methylations downstream of the repaired double-strand break.<a href="#cite_note-Cuozzo-108">[108]</a> Further work by Allen et al.<a href="#cite_note-pmid28423717-113">[113]</a> showed that NHEJ of a DNA double-strand break in a cell could give rise to some progeny cells having repressed expression of the gene harboring the initial double-strand break and some progeny having high expression of that gene due to epigenetic alterations associated with NHEJ repair. The frequency of epigenetic alterations causing repression of a gene after an NHEJ repair of a DNA double-strand break in that gene may be about 0.9%.<a href="#cite_note-pmid18704159-109">[109]</a> <div class="mw-heading mw-heading3"><h3 id="Techniques_used_to_study_epigenetics">Techniques used to study epigenetics</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=20" title="Edit section: Techniques used to study epigenetics">edit</a>]</div> Epigenetic research uses a wide range of <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biological</a> techniques to further understanding of epigenetic phenomena. These techniques include <a href="/wiki/Chromatin_immunoprecipitation" title="Chromatin immunoprecipitation">chromatin immunoprecipitation</a> (together with its large-scale variants <a href="/wiki/ChIP-on-chip" title="ChIP-on-chip">ChIP-on-chip</a> and <a href="/wiki/ChIP-Seq" class="mw-redirect" title="ChIP-Seq">ChIP-Seq</a>), <a href="/wiki/Fluorescent_in_situ_hybridization" class="mw-redirect" title="Fluorescent in situ hybridization">fluorescent in situ hybridization</a>, methylation-sensitive <a href="/wiki/Restriction_enzymes" class="mw-redirect" title="Restriction enzymes">restriction enzymes</a>, DNA adenine methyltransferase identification (<a href="/wiki/DamID" class="mw-redirect" title="DamID">DamID</a>) and <a href="/wiki/Bisulfite_sequencing" title="Bisulfite sequencing">bisulfite sequencing</a>.<a href="#cite_note-verma-114">[114]</a> Furthermore, the use of <a href="/wiki/Bioinformatics" title="Bioinformatics">bioinformatics</a> methods has a role in <a href="/wiki/Computational_epigenetics" title="Computational epigenetics">computational epigenetics</a>.<a href="#cite_note-verma-114">[114]</a> <div class="mw-heading mw-heading4"><h4 id="Chromatin_Immunoprecipitation">Chromatin Immunoprecipitation</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=21" title="Edit section: Chromatin Immunoprecipitation">edit</a>]</div> Chromatin Immunoprecipitation (ChIP) has helped bridge the gap between DNA and epigenetic interactions.<a href="#cite_note-Abcam-115">[115]</a> With the use of ChIP, researchers are able to make findings in regards to gene regulation, transcription mechanisms, and chromatin structure.<a href="#cite_note-Abcam-115">[115]</a> <div class="mw-heading mw-heading4"><h4 id="Fluorescent_in_situ_hybridization">Fluorescent in situ hybridization</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=22" title="Edit section: Fluorescent in situ hybridization">edit</a>]</div> Fluorescent in situ hybridization (FISH) is very important to understand epigenetic mechanisms.<a href="#cite_note-Chaumeil_2008-116">[116]</a> FISH can be used to find the location of genes on chromosomes, as well as finding noncoding RNAs.<a href="#cite_note-Chaumeil_2008-116">[116]</a><a href="#cite_note-O'Connor_2008-117">[117]</a> FISH is predominantly used for detecting chromosomal abnormalities in humans.<a href="#cite_note-O'Connor_2008-117">[117]</a> <div class="mw-heading mw-heading4"><h4 id="Methylation-sensitive_restriction_enzymes">Methylation-sensitive restriction enzymes</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=23" title="Edit section: Methylation-sensitive restriction enzymes">edit</a>]</div> Methylation sensitive restriction enzymes paired with PCR is a way to evaluate methylation in DNA - specifically the CpG sites.<a href="#cite_note-Hashimoto_2007-118">[118]</a> If DNA is methylated, the restriction enzymes will not cleave the strand.<a href="#cite_note-Hashimoto_2007-118">[118]</a> Contrarily, if the DNA is not methylated, the enzymes will cleave the strand and it will be amplified by PCR.<a href="#cite_note-Hashimoto_2007-118">[118]</a> <div class="mw-heading mw-heading4"><h4 id="Bisulfite_sequencing">Bisulfite sequencing</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=24" title="Edit section: Bisulfite sequencing">edit</a>]</div> Bisulfite sequencing is another way to evaluate DNA methylation. Cytosine will be changed to uracil from being treated with sodium bisulfite, whereas methylated cytosines will not be affected.<a href="#cite_note-Hashimoto_2007-118">[118]</a><a href="#cite_note-Li-Byarlay_et_al_2020-119">[119]</a><a href="#cite_note-ReferenceC-120">[120]</a> <div class="mw-heading mw-heading4"><h4 id="Nanopore_sequencing">Nanopore sequencing</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=25" title="Edit section: Nanopore sequencing">edit</a>]</div> Certain sequencing methods, such as <a href="/wiki/Nanopore_sequencing" title="Nanopore sequencing">nanopore sequencing</a>, allow sequencing of native DNA. Native (=unamplified) DNA retains the epigenetic modifications which would otherwise be lost during the amplification step. Nanopore basecaller models can distinguish between the signals obtained for epigenetically modified bases and unaltered based and provide an epigenetic profile in addition to the sequencing result.<a href="#cite_note-121">[121]</a> <div class="mw-heading mw-heading3"><h3 id="Structural_inheritance">Structural inheritance</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=26" title="Edit section: Structural inheritance">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/Structural_inheritance" title="Structural inheritance">Structural inheritance</a></div> In <a href="/wiki/Ciliate" title="Ciliate">ciliates</a> such as <a href="/wiki/Tetrahymena" title="Tetrahymena">Tetrahymena</a> and <a href="/wiki/Paramecium" title="Paramecium">Paramecium</a>, genetically identical cells show heritable differences in the patterns of ciliary rows on their cell surface. Experimentally altered patterns can be transmitted to daughter cells. It seems existing structures act as templates for new structures. The mechanisms of such inheritance are unclear, but reasons exist to assume that multicellular organisms also use existing cell structures to assemble new ones.<a href="#cite_note-pmid1804215-122">[122]</a><a href="#cite_note-isbn0-19-515619-6-123">[123]</a><a href="#cite_note-isbn0-262-65063-0-124">[124]</a> <div class="mw-heading mw-heading3"><h3 id="Nucleosome_positioning">Nucleosome positioning</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=27" title="Edit section: Nucleosome positioning">edit</a>]</div> Eukaryotic genomes have numerous <a href="/wiki/Nucleosomes" class="mw-redirect" title="Nucleosomes">nucleosomes</a>. Nucleosome position is not random, and determine the accessibility of DNA to regulatory proteins. Promoters active in different tissues have been shown to have different nucleosome positioning features.<a href="#cite_note-125">[125]</a> This determines differences in gene expression and cell differentiation. It has been shown that at least some nucleosomes are retained in sperm cells (where most but not all histones are replaced by <a href="/wiki/Protamines" class="mw-redirect" title="Protamines">protamines</a>). Thus nucleosome positioning is to some degree inheritable. Recent studies have uncovered connections between nucleosome positioning and other epigenetic factors, such as DNA methylation and hydroxymethylation.<a href="#cite_note-Teif_2014-126">[126]</a> <div class="mw-heading mw-heading3"><h3 id="Histone_variants">Histone variants</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=28" title="Edit section: Histone variants">edit</a>]</div> Different <a href="/wiki/Histone_variants" title="Histone variants">histone variants</a> are incorporated into specific regions of the genome non-randomly. Their differential biochemical characteristics can affect genome functions via their roles in gene regulation,<a href="#cite_note-127">[127]</a> and maintenance of chromosome structures.<a href="#cite_note-128">[128]</a> <div class="mw-heading mw-heading3"><h3 id="Genomic_architecture">Genomic architecture</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=29" title="Edit section: Genomic architecture">edit</a>]</div> The three-dimensional configuration of the genome (the 3D genome) is complex, dynamic and crucial for regulating genomic function and nuclear processes such as DNA replication, transcription and DNA-damage repair.<a href="#cite_note-129">[129]</a> <div class="mw-heading mw-heading2"><h2 id="Functions_and_consequences">Functions and consequences</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=30" title="Edit section: Functions and consequences">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="In_the_brain">In the brain</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=31" title="Edit section: In the brain">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="#Addiction">§ Addiction</a>, and <a href="#Depression">§ Depression</a></div> <div class="mw-heading mw-heading4"><h4 id="Memory">Memory</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=32" title="Edit section: Memory">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/Epigenetics_in_learning_and_memory" title="Epigenetics in learning and memory">Epigenetics in learning and memory</a></div> <a href="/wiki/Encoding_(memory)" title="Encoding (memory)">Memory formation</a> and maintenance are due to epigenetic alterations that cause the required dynamic changes in <a href="/wiki/Gene_transcription" class="mw-redirect" title="Gene transcription">gene transcription</a> that create and renew memory in neurons.<a href="#cite_note-Bernstein-31">[31]</a> An event can set off a chain of reactions that result in altered methylations of a large set of genes in neurons, which give a representation of the event, a memory.<a href="#cite_note-Bernstein-31">[31]</a> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Brain_regions_in_memory_formation_updated.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Brain_regions_in_memory_formation_updated.jpg/250px-Brain_regions_in_memory_formation_updated.jpg" decoding="async" width="250" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Brain_regions_in_memory_formation_updated.jpg/375px-Brain_regions_in_memory_formation_updated.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/58/Brain_regions_in_memory_formation_updated.jpg/500px-Brain_regions_in_memory_formation_updated.jpg 2x" data-file-width="1284" data-file-height="889" /></a><figcaption>including medial prefrontal cortex (mPFC)</figcaption></figure> Areas of the brain important in the formation of memories include the hippocampus, medial prefrontal cortex (mPFC), anterior cingulate cortex and amygdala, as shown in the diagram of the human brain in this section.<a href="#cite_note-pmid28386011-130">[130]</a> When a strong memory is created, as in a rat subjected to <a href="/wiki/Fear_conditioning" title="Fear conditioning">contextual fear conditioning</a> (CFC), one of the earliest events to occur is that more than 100 DNA double-strand breaks are formed by <a href="/wiki/Topoisomerase" title="Topoisomerase">topoisomerase IIB</a> in neurons of the hippocampus and the medial prefrontal cortex (mPFC).<a href="#cite_note-Stott-131">[131]</a> These double-strand breaks are at specific locations that allow activation of transcription of <a href="/wiki/Immediate_early_genes" class="mw-redirect" title="Immediate early genes">immediate early genes</a> (IEGs) that are important in memory formation, allowing their expression in <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a>, with peak mRNA transcription at seven to ten minutes after CFC.<a href="#cite_note-Stott-131">[131]</a><a href="#cite_note-pmid35776545-132">[132]</a> Two important IEGs in memory formation are <a href="/wiki/EGR1" title="EGR1">EGR1</a><a href="#cite_note-pmid10357227-133">[133]</a> and <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">the alternative promoter variant of DNMT3A, DNMT3A2</a>.<a href="#cite_note-pmid22751036-134">[134]</a> EGR1 protein binds to DNA at its binding motifs, 5′-GCGTGGGCG-3′ or 5′-GCGGGGGCGG-3', and there are about 12,000 genome locations at which EGR1 protein can bind.<a href="#cite_note-Sun-135">[135]</a> EGR1 protein binds to DNA in gene <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions. EGR1 recruits the demethylating enzyme <a href="/wiki/TET_enzymes" title="TET enzymes">TET1</a> to an association, and brings TET1 to about 600 locations on the genome where TET1 can then demethylate and activate the associated genes.<a href="#cite_note-Sun-135">[135]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cytosine_and_5-methylcytosine.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Cytosine_and_5-methylcytosine.jpg/220px-Cytosine_and_5-methylcytosine.jpg" decoding="async" width="220" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Cytosine_and_5-methylcytosine.jpg/330px-Cytosine_and_5-methylcytosine.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Cytosine_and_5-methylcytosine.jpg/440px-Cytosine_and_5-methylcytosine.jpg 2x" data-file-width="1524" data-file-height="954" /></a><figcaption>Cytosine and 5-methylcytosine</figcaption></figure> The DNA methyltransferases DNMT3A1, DNMT3A2 and DNMT3B can all methylate cytosines (see image this section) at <a href="/wiki/CpG_site" title="CpG site">CpG sites</a> in or near the promoters of genes. As shown by Manzo et al.,<a href="#cite_note-pmid29074627-136">[136]</a> these three DNA methyltransferases differ in their genomic binding locations and DNA methylation activity at different regulatory sites. Manzo et al. located 3,970 genome regions exclusively enriched for DNMT3A1, 3,838 regions for DNMT3A2 and 3,432 regions for DNMT3B. When DNMT3A2 is newly induced as an IEG (when neurons are activated), many new cytosine methylations occur, presumably in the target regions of DNMT3A2. Oliviera et al.<a href="#cite_note-pmid22751036-134">[134]</a> found that the neuronal activity-inducible IEG levels of Dnmt3a2 in the hippocampus determined the ability to form long-term memories. Rats form long-term associative memories after <a href="/wiki/Fear_conditioning" title="Fear conditioning">contextual fear conditioning (CFC)</a>.<a href="#cite_note-pmid25324744-137">[137]</a> Duke et al.<a href="#cite_note-pmid28620075-30">[30]</a> found that 24 hours after CFC in rats, in hippocampus neurons, 2,097 genes (9.17% of the genes in the rat genome) had altered methylation. When newly methylated cytosines are present in <a href="/wiki/CpG_site" title="CpG site">CpG sites</a> in the promoter regions of genes, the genes are often repressed, and when newly demethylated cytosines are present the genes may be activated.<a href="#cite_note-pmid22781841-138">[138]</a> After CFC, there were 1,048 genes with reduced mRNA expression and 564 genes with upregulated mRNA expression. Similarly, when mice undergo CFC, one hour later in the hippocampus region of the mouse brain there are 675 demethylated genes and 613 hypermethylated genes.<a href="#cite_note-Halder-139">[139]</a> However, memories do not remain in the hippocampus, but after four or five weeks the memories are stored in the anterior cingulate cortex.<a href="#cite_note-pmid15131309-140">[140]</a> In the studies on mice after CFC, Halder et al.<a href="#cite_note-Halder-139">[139]</a> showed that four weeks after CFC there were at least 1,000 differentially methylated genes and more than 1,000 differentially expressed genes in the anterior cingulate cortex, while at the same time the altered methylations in the hippocampus were reversed. The epigenetic alteration of methylation after a new memory is established creates a different pool of nuclear mRNAs. As reviewed by Bernstein,<a href="#cite_note-Bernstein-31">[31]</a> the epigenetically determined new mix of nuclear <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNAs</a> are often packaged into neuronal granules, or <a href="/wiki/Messenger_RNP" title="Messenger RNP">messenger RNP</a>, consisting of mRNA, <a href="/wiki/Ribosome" title="Ribosome">small and large ribosomal subunits</a>, translation initiation factors and RNA-binding proteins that regulate mRNA function. These neuronal granules are transported from the neuron nucleus and are directed, according to 3′ untranslated regions of the mRNA in the granules (their "zip codes"), to neuronal <a href="/wiki/Dendrite" title="Dendrite">dendrites</a>. Roughly 2,500 mRNAs may be localized to the dendrites of hippocampal pyramidal neurons and perhaps 450 transcripts are in excitatory presynaptic nerve terminals (dendritic spines). The altered assortments of transcripts (dependent on epigenetic alterations in the neuron nucleus) have different sensitivities in response to signals, which is the basis of altered synaptic plasticity. Altered synaptic plasticity is often considered the neurochemical foundation of learning and memory. <div class="mw-heading mw-heading4"><h4 id="Aging">Aging</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=33" title="Edit section: Aging">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/DNA_methylation#In_aging" title="DNA methylation">DNA methylation § In aging</a>, and <a href="/wiki/Hallmarks_of_aging#Epigenomic_alterations" title="Hallmarks of aging">Hallmarks of aging § Epigenomic alterations</a></div> Epigenetics play a major role in <a href="/wiki/Brain_aging" class="mw-redirect" title="Brain aging">brain aging</a> and age-related cognitive decline, with relevance to <a href="/wiki/Life_extension" title="Life extension">life extension</a>.<a href="#cite_note-141">[141]</a><a href="#cite_note-142">[142]</a><a href="#cite_note-143">[143]</a><a href="#cite_note-144">[144]</a><a href="#cite_note-145">[145]</a> <div class="mw-heading mw-heading4"><h4 id="Other_and_general">Other and general</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=34" title="Edit section: Other and general">edit</a>]</div> In adulthood, changes in the <a href="/wiki/Epigenome" title="Epigenome">epigenome</a> are important for various higher cognitive functions. Dysregulation of epigenetic mechanisms is implicated in <a href="/wiki/Neurodegenerative_disorders" class="mw-redirect" title="Neurodegenerative disorders">neurodegenerative disorders</a> and diseases. Epigenetic modifications in <a href="/wiki/Neuron" title="Neuron">neurons</a> are dynamic and reversible.<a href="#cite_note-146">[146]</a> Epigenetic regulation impacts neuronal action, affecting learning, memory, and other <a href="/wiki/Cognitive" class="mw-redirect" title="Cognitive">cognitive</a> processes.<a href="#cite_note-147">[147]</a> Early events, including during <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a>, can influence development, cognition, and health outcomes through <a href="/w/index.php?title=Epigenetic_mechanisms&action=edit&redlink=1" class="new" title="Epigenetic mechanisms (page does not exist)">epigenetic mechanisms</a>.<a href="#cite_note-148">[148]</a> Epigenetic mechanisms have been proposed as "a potential molecular mechanism for effects of endogenous <a href="/wiki/Hormone" title="Hormone">hormones</a> on the organization of developing brain circuits".<a href="#cite_note-149">[149]</a> <a href="/wiki/Nutrients" class="mw-redirect" title="Nutrients">Nutrients</a> could interact with the epigenome to "protect or boost cognitive processes across the lifespan".<a href="#cite_note-150">[150]</a><a href="#cite_note-151">[151]</a> A review suggests <a href="/wiki/Neurobiological_effects_of_physical_exercise" title="Neurobiological effects of physical exercise">neurobiological effects of physical exercise</a> via <a href="/wiki/Epigenetics_of_physical_exercise" title="Epigenetics of physical exercise">epigenetics</a> seem "central to building an 'epigenetic memory' to influence long-term brain function and behavior" and may even be heritable.<a href="#cite_note-152">[152]</a> With the axo-ciliary <a href="/wiki/Synapse" title="Synapse">synapse</a>, there is communication between <a href="/wiki/Serotonin" title="Serotonin">serotonergic</a> <a href="/wiki/Axon" title="Axon">axons</a> and antenna-like <a href="/wiki/Primary_cilia" class="mw-redirect" title="Primary cilia">primary cilia</a> of <a href="/wiki/Hippocampus_anatomy#Basic_hippocampal_circuit" title="Hippocampus anatomy">CA1</a> <a href="/wiki/Pyramidal_cell" title="Pyramidal cell">pyramidal</a> <a href="/wiki/Neuron" title="Neuron">neurons</a> that alters the neuron's <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> state in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> via the signalling distinct from that at the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> (and longer-term).<a href="#cite_note-153">[153]</a><a href="#cite_note-154">[154]</a> Epigenetics also play a major role in the <a href="/wiki/Evolution_of_the_brain#Genetic_factors_of_recent_evolution" title="Evolution of the brain">brain evolution in and to humans</a>.<a href="#cite_note-155">[155]</a> <div class="mw-heading mw-heading3"><h3 id="Development">Development</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=35" title="Edit section: Development">edit</a>]</div> Developmental epigenetics can be divided into predetermined and probabilistic epigenesis. Predetermined epigenesis is a unidirectional movement from structural development in DNA to the functional maturation of the protein. "Predetermined" here means that development is scripted and predictable. Probabilistic epigenesis on the other hand is a bidirectional structure-function development with experiences and external molding development.<a href="#cite_note-Griesemer_2005-156">[156]</a> Somatic epigenetic inheritance, particularly through DNA and histone covalent modifications and <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a> repositioning, is very important in the development of multicellular eukaryotic organisms.<a href="#cite_note-Teif_2014-126">[126]</a> The genome sequence is static (with some notable exceptions), but cells differentiate into many different types, which perform different functions, and respond differently to the environment and intercellular signaling. Thus, as individuals develop, <a href="/wiki/Morphogen" title="Morphogen">morphogens</a> activate or silence genes in an epigenetically heritable fashion, giving cells a memory. In mammals, most cells terminally differentiate, with only <a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">stem cells</a> retaining the ability to differentiate into several cell types ("totipotency" and "multipotency"). In <a href="/wiki/Mammal" title="Mammal">mammals</a>, some stem cells continue producing newly differentiated cells throughout life, such as in <a href="/wiki/Epigenetic_Regulation_of_Neurogenesis" class="mw-redirect" title="Epigenetic Regulation of Neurogenesis">neurogenesis</a>, but mammals are not able to respond to loss of some tissues, for example, the inability to regenerate limbs, which some other animals are capable of. Epigenetic modifications regulate the transition from neural stem cells to glial progenitor cells (for example, differentiation into oligodendrocytes is regulated by the deacetylation and methylation of histones).<a href="#cite_note-157">[157]</a> Unlike animals, plant cells do not terminally differentiate, remaining totipotent with the ability to give rise to a new individual plant. While plants do utilize many of the same epigenetic mechanisms as animals, such as <a href="/wiki/Chromatin_remodeling" title="Chromatin remodeling">chromatin remodeling</a>, it has been hypothesized that some kinds of plant cells do not use or require "cellular memories", resetting their gene expression patterns using positional information from the environment and surrounding cells to determine their fate.<a href="#cite_note-pmid17194589-158">[158]</a> Epigenetic changes can occur in response to environmental exposure – for example, maternal dietary supplementation with <a href="/wiki/Genistein" title="Genistein">genistein</a> (250 mg/kg) have epigenetic changes affecting expression of the <a href="/wiki/Agouti_gene" class="mw-redirect" title="Agouti gene">agouti gene</a>, which affects their fur color, weight, and propensity to develop cancer.<a href="#cite_note-pmid12163699-159">[159]</a><a href="#cite_note-waterland-160">[160]</a><a href="#cite_note-161">[161]</a> Ongoing research is focused on exploring the impact of other known <a href="/wiki/Teratogen" class="mw-redirect" title="Teratogen">teratogens</a>, such as <a href="/wiki/Diabetic_embryopathy" title="Diabetic embryopathy">diabetic embryopathy</a>, on <a href="/wiki/Methylation" title="Methylation">methylation</a> signatures.<a href="#cite_note-162">[162]</a> Controversial results from one study suggested that traumatic experiences might produce an epigenetic signal that is capable of being passed to future generations. Mice were trained, using foot shocks, to fear a cherry blossom odor. The investigators reported that the mouse offspring had an increased aversion to this specific odor.<a href="#cite_note-163">[163]</a><a href="#cite_note-164">[164]</a> They suggested epigenetic changes that increase gene expression, rather than in DNA itself, in a gene, M71, that governs the functioning of an odor receptor in the nose that responds specifically to this cherry blossom smell. There were physical changes that correlated with olfactory (smell) function in the brains of the trained mice and their descendants. Several criticisms were reported, including the study's low statistical power as evidence of some irregularity such as bias in reporting results.<a href="#cite_note-Francis_2014-165">[165]</a> Due to limits of sample size, there is a probability that an effect will not be demonstrated to within statistical significance even if it exists. The criticism suggested that the probability that all the experiments reported would show positive results if an identical protocol was followed, assuming the claimed effects exist, is merely 0.4%. The authors also did not indicate which mice were siblings, and treated all of the mice as statistically independent.<a href="#cite_note-166">[166]</a> The original researchers pointed out negative results in the paper's appendix that the criticism omitted in its calculations, and undertook to track which mice were siblings in the future.<a href="#cite_note-167">[167]</a> <div class="mw-heading mw-heading3"><h3 id="Transgenerational">Transgenerational</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=36" title="Edit section: Transgenerational">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/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></div> Epigenetic mechanisms were a necessary part of the evolutionary origin of <a href="/wiki/Cell_differentiation" class="mw-redirect" title="Cell differentiation">cell differentiation</a>.<a href="#cite_note-isbn0-19-854968-7-168">[168]</a>[<a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">need quotation to verify</a>] Although epigenetics in multicellular organisms is generally thought to be a mechanism involved in differentiation, with epigenetic patterns "reset" when organisms reproduce, there have been some observations of transgenerational epigenetic inheritance (e.g., the phenomenon of <a href="/wiki/Paramutation" title="Paramutation">paramutation</a> observed in <a href="/wiki/Maize" title="Maize">maize</a>). Although most of these multigenerational epigenetic traits are gradually lost over several generations, the possibility remains that multigenerational epigenetics could be another aspect to <a href="/wiki/Evolution" title="Evolution">evolution</a> and adaptation. As mentioned above, some define epigenetics as heritable. A sequestered germ line or <a href="/wiki/Weismann_barrier" title="Weismann barrier">Weismann barrier</a> is specific to animals, and epigenetic inheritance is more common in plants and microbes. <a href="/wiki/Eva_Jablonka" title="Eva Jablonka">Eva Jablonka</a>, <a href="/wiki/Marion_J._Lamb" title="Marion J. Lamb">Marion J. Lamb</a> and Étienne Danchin have argued that these effects may require enhancements to the standard conceptual framework of the <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern synthesis</a> and have called for an <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">extended evolutionary synthesis</a>.<a href="#cite_note-isbn0-262-10107-6-169">[169]</a><a href="#cite_note-170">[170]</a><a href="#cite_note-171">[171]</a> Other evolutionary biologists, such as <a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">John Maynard Smith</a>, have incorporated epigenetic inheritance into <a href="/wiki/Population_genetics" title="Population genetics">population-genetics</a> models<a href="#cite_note-172">[172]</a> or are openly skeptical of the extended evolutionary synthesis (<a href="/wiki/Michael_Lynch_(geneticist)" title="Michael Lynch (geneticist)">Michael Lynch</a>).<a href="#cite_note-173">[173]</a> Thomas Dickins and Qazi Rahman state that epigenetic mechanisms such as DNA methylation and histone modification are genetically inherited under the control of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> and therefore fit under the earlier <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">"modern synthesis"</a>.<a href="#cite_note-174">[174]</a> Two important ways in which epigenetic inheritance can differ from traditional genetic inheritance, with important consequences for evolution, are: <ul><li>rates of epimutation can be much faster than rates of mutation<a href="#cite_note-rando_and_verstrepen-175">[175]</a></li> <li>the epimutations are more easily reversible<a href="#cite_note-176">[176]</a></li></ul> In plants, heritable DNA methylation mutations are 100,000 times more likely to occur compared to DNA mutations.<a href="#cite_note-van_der_Graaf_et_al-177">[177]</a> An epigenetically inherited element such as the <a href="/wiki/PSI_(prion)" class="mw-redirect" title="PSI (prion)">PSI+</a> system can act as a "stop-gap", good enough for short-term adaptation that allows the lineage to survive for long enough for mutation and/or recombination to <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetically assimilate</a> the adaptive phenotypic change.<a href="#cite_note-178">[178]</a> The existence of this possibility increases the <a href="/wiki/Evolvability" title="Evolvability">evolvability</a> of a species. More than 100 cases of <a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">transgenerational epigenetic inheritance</a> phenomena have been reported in a wide range of organisms, including prokaryotes, plants, and animals.<a href="#cite_note-Jablonka09-179">[179]</a> For instance, <a href="/wiki/Nymphalis_antiopa" title="Nymphalis antiopa">mourning-cloak butterflies</a> will change color through hormone changes in response to experimentation of varying temperatures.<a href="#cite_note-180">[180]</a> The filamentous fungus Neurospora crassa is a prominent model system for understanding the control and function of cytosine methylation. In this organism, DNA methylation is associated with relics of a genome-defense system called RIP (repeat-induced point mutation) and silences gene expression by inhibiting transcription elongation.<a href="#cite_note-pmid19092133-181">[181]</a> The <a href="/wiki/Yeast_prion" class="mw-redirect" title="Yeast prion">yeast prion</a> PSI is generated by a conformational change of a translation termination factor, which is then inherited by daughter cells. This can provide a survival advantage under adverse conditions, exemplifying epigenetic regulation which enables unicellular organisms to respond rapidly to environmental stress. Prions can be viewed as epigenetic agents capable of inducing a phenotypic change without modification of the genome.<a href="#cite_note-JorgTost-182">[182]</a> Direct detection of epigenetic marks in microorganisms is possible with <a href="/wiki/Single_molecule_real_time_sequencing" class="mw-redirect" title="Single molecule real time sequencing">single molecule real time sequencing</a>, in which polymerase sensitivity allows for measuring methylation and other modifications as a DNA molecule is being sequenced.<a href="#cite_note-183">[183]</a> Several projects have demonstrated the ability to collect genome-wide epigenetic data in bacteria.<a href="#cite_note-184">[184]</a><a href="#cite_note-185">[185]</a><a href="#cite_note-186">[186]</a><a href="#cite_note-187">[187]</a> <div class="mw-heading mw-heading2"><h2 id="Epigenetics_in_bacteria">Epigenetics in bacteria</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=37" title="Edit section: Epigenetics in bacteria">edit</a>]</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Escherichia_coli_flagella_TEM.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Escherichia_coli_flagella_TEM.png/150px-Escherichia_coli_flagella_TEM.png" decoding="async" width="150" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Escherichia_coli_flagella_TEM.png/225px-Escherichia_coli_flagella_TEM.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Escherichia_coli_flagella_TEM.png/300px-Escherichia_coli_flagella_TEM.png 2x" data-file-width="600" data-file-height="896" /></a><figcaption>Escherichia coli bacteria</figcaption></figure> While epigenetics is of fundamental importance in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, especially <a href="/wiki/Multicellular_organism" title="Multicellular organism">metazoans</a>, it plays a different role in bacteria.<a href="#cite_note-188">[188]</a> Most importantly, eukaryotes use epigenetic mechanisms primarily to regulate gene expression which bacteria rarely do. However, bacteria make widespread use of postreplicative DNA methylation for the epigenetic control of DNA-protein interactions. Bacteria also use DNA <a href="/wiki/Adenine" title="Adenine">adenine</a> methylation (rather than DNA <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> methylation) as an epigenetic signal. DNA adenine methylation is important in bacteria virulence in organisms such as <a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a>, <a href="/wiki/Salmonella" title="Salmonella">Salmonella</a>, <a href="/wiki/Vibrio" title="Vibrio">Vibrio</a>, <a href="/wiki/Yersinia" title="Yersinia">Yersinia</a>, <a href="/wiki/Haemophilus" title="Haemophilus">Haemophilus</a>, and <a href="/wiki/Brucella" title="Brucella">Brucella</a>. In <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">Alphaproteobacteria</a>, methylation of adenine regulates the cell cycle and couples gene transcription to DNA replication. In <a href="/wiki/Gammaproteobacteria" title="Gammaproteobacteria">Gammaproteobacteria</a>, adenine methylation provides signals for DNA replication, chromosome segregation, mismatch repair, packaging of bacteriophage, transposase activity and regulation of gene expression.<a href="#cite_note-JorgTost-182">[182]</a><a href="#cite_note-Casadesus-189">[189]</a> There exists a genetic switch controlling <a href="/wiki/Streptococcus_pneumoniae" title="Streptococcus pneumoniae">Streptococcus pneumoniae</a> (the pneumococcus) that allows the bacterium to randomly change its characteristics into six alternative states that could pave the way to improved vaccines. Each form is randomly generated by a phase variable methylation system. The ability of the pneumococcus to cause deadly infections is different in each of these six states. Similar systems exist in other bacterial genera.<a href="#cite_note-MansoOggioni2014-190">[190]</a> In <a href="/wiki/Bacillota" title="Bacillota">Bacillota</a> such as <a href="/wiki/Clostridioides_difficile_(bacteria)" class="mw-redirect" title="Clostridioides difficile (bacteria)">Clostridioides difficile</a>, adenine methylation regulates <a href="/wiki/Spore" title="Spore">sporulation</a>, <a href="/wiki/Biofilm" title="Biofilm">biofilm</a> formation and host-adaptation.<a href="#cite_note-191">[191]</a> <div class="mw-heading mw-heading2"><h2 id="Medicine">Medicine</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=38" title="Edit section: Medicine">edit</a>]</div> Epigenetics has many and varied potential medical applications.<a href="#cite_note-pmid21447282-192">[192]</a> <div class="mw-heading mw-heading3"><h3 id="Twins">Twins</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=39" title="Edit section: Twins">edit</a>]</div> Direct comparisons of identical twins constitute an optimal model for interrogating <a href="/wiki/Environmental_epigenetics" title="Environmental epigenetics">environmental epigenetics</a>. In the case of humans with different environmental exposures, monozygotic (identical) twins were epigenetically indistinguishable during their early years, while older twins had remarkable differences in the overall content and genomic distribution of 5-methylcytosine DNA and histone acetylation.<a href="#cite_note-Moore_2015-11">[11]</a> The twin pairs who had spent less of their lifetime together and/or had greater differences in their medical histories were those who showed the largest differences in their levels of <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a> DNA and <a href="/wiki/Acetylation" title="Acetylation">acetylation</a> of <a href="/wiki/Histones" class="mw-redirect" title="Histones">histones</a> H3 and H4.<a href="#cite_note-pmid16009939-193">[193]</a> Dizygotic (fraternal) and monozygotic (identical) twins show evidence of epigenetic influence in humans.<a href="#cite_note-pmid16009939-193">[193]</a><a href="#cite_note-pmid19151718-194">[194]</a><a href="#cite_note-195">[195]</a> DNA sequence differences that would be abundant in a singleton-based study do not interfere with the analysis. Environmental differences can produce long-term epigenetic effects, and different developmental monozygotic twin subtypes may be different with respect to their susceptibility to be discordant from an epigenetic point of view.<a href="#cite_note-pmid19653134-196">[196]</a> A <a href="/wiki/High-throughput_screening" title="High-throughput screening">high-throughput</a> study, which denotes technology that looks at extensive genetic markers, focused on epigenetic differences between monozygotic twins to compare global and locus-specific changes in <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> and histone modifications in a sample of 40 monozygotic twin pairs.<a href="#cite_note-pmid16009939-193">[193]</a> In this case, only healthy twin pairs were studied, but a wide range of ages was represented, between 3 and 74 years. One of the major conclusions from this study was that there is an age-dependent accumulation of epigenetic differences between the two siblings of twin pairs. This accumulation suggests the existence of epigenetic "drift". Epigenetic drift is the term given to epigenetic modifications as they occur as a direct function with age. While age is a known risk factor for many diseases, age-related methylation has been found to occur differentially at specific sites along the genome. Over time, this can result in measurable differences between biological and chronological age. Epigenetic changes have been found to be reflective of <a href="/wiki/Lifestyle_(social_sciences)" title="Lifestyle (social sciences)">lifestyle</a> and may act as functional <a href="/wiki/Biomarker" title="Biomarker">biomarkers</a> of disease before clinical <a href="/wiki/Reference_range" title="Reference range">threshold</a> is reached.<a href="#cite_note-197">[197]</a> A more recent study, where 114 monozygotic twins and 80 dizygotic twins were analyzed for the DNA methylation status of around 6000 unique genomic regions, concluded that epigenetic similarity at the time of blastocyst splitting may also contribute to phenotypic similarities in monozygotic co-twins. This supports the notion that <a href="/wiki/Microenvironment_(biology)" class="mw-redirect" title="Microenvironment (biology)">microenvironment</a> at early stages of embryonic development can be quite important for the establishment of epigenetic marks.<a href="#cite_note-pmid19151718-194">[194]</a> Congenital genetic disease is well understood and it is clear that epigenetics can play a role, for example, in the case of <a href="/wiki/Angelman_syndrome" title="Angelman syndrome">Angelman syndrome</a> and <a href="/wiki/Prader%E2%80%93Willi_syndrome" title="Prader–Willi syndrome">Prader–Willi syndrome</a>. These are normal genetic diseases caused by gene deletions or inactivation of the genes but are unusually common because individuals are essentially <a href="/wiki/Hemizygous" class="mw-redirect" title="Hemizygous">hemizygous</a> because of <a href="/wiki/Genomic_imprinting" title="Genomic imprinting">genomic imprinting</a>, and therefore a single gene knock out is sufficient to cause the disease, where most cases would require both copies to be knocked out.<a href="#cite_note-198">[198]</a> <div class="mw-heading mw-heading3"><h3 id="Genomic_imprinting">Genomic imprinting</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=40" title="Edit section: Genomic imprinting">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/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></div> Some human disorders are associated with genomic imprinting, a phenomenon in mammals where the father and mother contribute different epigenetic patterns for specific genomic loci in their <a href="/wiki/Germ_cells" class="mw-redirect" title="Germ cells">germ cells</a>.<a href="#cite_note-pmid17121465-199">[199]</a> The best-known case of imprinting in human disorders is that of <a href="/wiki/Angelman_syndrome" title="Angelman syndrome">Angelman syndrome</a> and <a href="/wiki/Prader%E2%80%93Willi_syndrome" title="Prader–Willi syndrome">Prader–Willi syndrome</a> – both can be produced by the same genetic mutation, <a href="/wiki/Chromosome_15q_partial_deletion" title="Chromosome 15q partial deletion">chromosome 15q partial deletion</a>, and the particular syndrome that will develop depends on whether the mutation is inherited from the child's mother or from their father.<a href="#cite_note-pmid2564739-200">[200]</a> In the <a href="/wiki/%C3%96verkalix_study" title="Överkalix study">Överkalix study</a>, paternal (but not maternal) grandsons<a href="#cite_note-paternal-grandson-201">[201]</a> of Swedish men who were exposed during preadolescence to famine in the 19th century were less likely to die of cardiovascular disease. If food was plentiful, then <a href="/wiki/Diabetes" title="Diabetes">diabetes</a> mortality in the grandchildren increased, suggesting that this was a transgenerational epigenetic inheritance.<a href="#cite_note-pmid16391557-202">[202]</a> The opposite effect was observed for females – the paternal (but not maternal) granddaughters of women who experienced famine while in the womb (and therefore while their eggs were being formed) lived shorter lives on average.<a href="#cite_note-203">[203]</a> <div class="mw-heading mw-heading3"><h3 id="Examples_of_drugs_altering_gene_expression_from_epigenetic_events">Examples of drugs altering gene expression from epigenetic events</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=41" title="Edit section: Examples of drugs altering gene expression from epigenetic events">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Epigenetic_Priming" class="mw-redirect" title="Epigenetic Priming">Epigenetic Priming</a></div> The use of beta-lactam <a href="/wiki/Antibiotics" class="mw-redirect" title="Antibiotics">antibiotics</a> can alter glutamate receptor activity and the action of cyclosporine on multiple transcription factors. Additionally, <a href="/wiki/Lithium" title="Lithium">lithium</a> can impact autophagy of aberrant proteins, and <a href="/wiki/Opioid" title="Opioid">opioid</a> drugs via chronic use can increase the expression of genes associated with addictive phenotypes.<a href="#cite_note-204">[204]</a> Parental <a href="/wiki/Nutrition" title="Nutrition">nutrition</a>, in utero exposure to stress or <a href="/wiki/Endocrine_disruptor" title="Endocrine disruptor">endocrine disrupting chemicals</a>,<a href="#cite_note-205">[205]</a> male-induced maternal effects such as the attraction of differential mate quality, and maternal as well as paternal age, and offspring gender could all possibly influence whether a germline epimutation is ultimately expressed in offspring and the degree to which intergenerational inheritance remains stable throughout posterity.<a href="#cite_note-ReferenceB-206">[206]</a> However, whether and to what extent epigenetic effects can be transmitted across generations remains unclear, particularly in humans.<a href="#cite_note-PlominDeFries2012-207">[207]</a><a href="#cite_note-208">[208]</a> <div class="mw-heading mw-heading3"><h3 id="Addiction">Addiction</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=42" title="Edit section: Addiction">edit</a>]</div> <a href="/wiki/Addiction" title="Addiction">Addiction</a> is a disorder of the brain's <a href="/wiki/Reward_system" title="Reward system">reward system</a> which arises through <a href="/wiki/Transcriptional" class="mw-redirect" title="Transcriptional">transcriptional</a> and neuroepigenetic mechanisms and occurs over time from chronically high levels of exposure to an addictive stimulus (e.g., morphine, cocaine, sexual intercourse, gambling).<a href="#cite_note-Nestler-209">[209]</a><a href="#cite_note-Cheron2021-210">[210]</a><a href="#cite_note-G9a_reverses_ΔFosB_plasticity-211">[211]</a> Transgenerational epigenetic inheritance of addictive <a href="/wiki/Phenotypes" class="mw-redirect" title="Phenotypes">phenotypes</a> has been noted to occur in preclinical studies.<a href="#cite_note-pmid23920159-212">[212]</a><a href="#cite_note-pmid26572641-213">[213]</a> However, robust evidence in support of the persistence of epigenetic effects across multiple generations has yet to be established in humans; for example, an epigenetic effect of prenatal exposure to smoking that is observed in great-grandchildren who had not been exposed.<a href="#cite_note-PlominDeFries2012-207">[207]</a> <div class="mw-heading mw-heading2"><h2 id="Research">Research</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=43" title="Edit section: Research">edit</a>]</div> The two forms of heritable information, namely genetic and epigenetic, are collectively called dual inheritance. Members of the APOBEC/AID family of <a href="/wiki/Cytosine_deaminase" title="Cytosine deaminase">cytosine deaminases</a> may concurrently influence genetic and epigenetic inheritance using similar molecular mechanisms, and may be a point of crosstalk between these conceptually compartmentalized processes.<a href="#cite_note-pmid20800313-214">[214]</a> <a href="/wiki/Fluoroquinolone" class="mw-redirect" title="Fluoroquinolone">Fluoroquinolone</a> antibiotics induce epigenetic changes in <a href="/wiki/Mammalian" class="mw-redirect" title="Mammalian">mammalian</a> cells through iron <a href="/wiki/Chelation" title="Chelation">chelation</a>. This leads to epigenetic effects through inhibition of α-ketoglutarate-dependent <a href="/wiki/Dioxygenases" class="mw-redirect" title="Dioxygenases">dioxygenases</a> that require <a href="/wiki/Iron" title="Iron">iron</a> as a co-factor.<a href="#cite_note-215">[215]</a> Various pharmacological agents are applied for the production of induced pluripotent stem cells (iPSC) or maintain the embryonic stem cell (ESC) phenotypic via epigenetic approach. Adult stem cells like bone marrow stem cells have also shown a potential to differentiate into cardiac competent cells when treated with G9a histone methyltransferase inhibitor BIX01294.<a href="#cite_note-216">[216]</a><a href="#cite_note-217">[217]</a> Cell plasticity, which is the adaptation of cells to stimuli without changes in their genetic code, requires epigenetic changes. These have been observed in cell plasticity in cancer cells during epithelial-to-mesenchymal transition<a href="#cite_note-218">[218]</a> and also in immune cells, such as macrophages.<a href="#cite_note-219">[219]</a> Interestingly, metabolic changes underly these adaptations, since various metabolites play crucial roles in the chemistry of epigenetic marks. This includes for instance alpha-ketoglutarate, which is required for histone demethylation, and acetyl-Coenzyme A, which is required for histone acetylation. <div class="mw-heading mw-heading3"><h3 id="Epigenome_editing">Epigenome editing</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=44" title="Edit section: Epigenome editing">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/Epigenome_editing" title="Epigenome editing">Epigenome editing</a></div> Epigenetic regulation of gene expression that could be altered or used in <a href="/wiki/Epigenome_editing" title="Epigenome editing">epigenome editing</a> are or include <a href="/w/index.php?title=MRNA_modification&action=edit&redlink=1" class="new" title="MRNA modification (page does not exist)">mRNA/lncRNA modification</a>, <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> modification and <a href="/wiki/Histone_modification" class="mw-redirect" title="Histone modification">histone modification</a>.<a href="#cite_note-220">[220]</a><a href="#cite_note-221">[221]</a><a href="#cite_note-222">[222]</a> <div class="mw-heading mw-heading3"><h3 id="CpG_sites,_SNPs_and_biological_traits">CpG sites, SNPs and biological traits</h3>[<a href="/w/index.php?title=Epigenetics&action=edit&section=45" title="Edit section: CpG sites, SNPs and biological traits">edit</a>]</div> Methylation is a widely characterized mechanism of genetic regulation that can determine biological traits. However, strong experimental evidences correlate methylation patterns in SNPs as an important additional feature for the classical activation/inhibition epigenetic dogma. Molecular interaction data, supported by colocalization analyses, identify multiple nuclear regulatory pathways, linking sequence variation to disturbances in DNA methylation and molecular and phenotypic variation.<a href="#cite_note-Hawe_2022-223">[223]</a> <div class="mw-heading mw-heading4"><h4 id="UBASH3B_locus">UBASH3B locus</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=46" title="Edit section: UBASH3B locus">edit</a>]</div> UBASH3B encodes a protein with tyrosine phosphatase activity, which has been previously linked to advanced neoplasia.<a href="#cite_note-224">[224]</a> SNP rs7115089 was identified as influencing DNA methylation and expression of this locus, as well as and Body Mass Index (BMI).<a href="#cite_note-Hawe_2022-223">[223]</a> In fact, SNP rs7115089 is strongly associated with BMI<a href="#cite_note-225">[225]</a> and with genetic variants linked to other cardiovascular and metabolic traits in GWASs.<a href="#cite_note-226">[226]</a><a href="#cite_note-227">[227]</a><a href="#cite_note-228">[228]</a> New studies suggesting UBASH3B as a potential mediator of adiposity and cardiometabolic disease.<a href="#cite_note-Hawe_2022-223">[223]</a> In addition, animal models demonstrated that UBASH3B expression is an indicator of caloric restriction that may drive programmed susceptibility to obesity and it is associated with other measures of adiposity in human peripherical blood.<a href="#cite_note-229">[229]</a> <div class="mw-heading mw-heading4"><h4 id="NFKBIE_locus">NFKBIE locus</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=47" title="Edit section: NFKBIE locus">edit</a>]</div> SNP rs730775 is located in the first intron of NFKBIE and is a cis eQTL for NFKBIE in whole blood.<a href="#cite_note-Hawe_2022-223">[223]</a> Nuclear factor (NF)-κB inhibitor ε (NFKBIE) directly inhibits NF-κB1 activity and is significantly co-expressed with NF-κB1, also, it is associated with rheumatoid arthritis.<a href="#cite_note-230">[230]</a> Colocalization analysis supports that variants for the majority of the CpG sites in SNP rs730775 cause genetic variation at the NFKBIE locus which is suggestible linked to rheumatoid arthritis through trans acting regulation of DNA methylation by NF-κB.<a href="#cite_note-Hawe_2022-223">[223]</a> <div class="mw-heading mw-heading4"><h4 id="FADS1_locus">FADS1 locus</h4>[<a href="/w/index.php?title=Epigenetics&action=edit&section=48" title="Edit section: FADS1 locus">edit</a>]</div> Fatty acid desaturase 1 (FADS1) is a key enzyme in the metabolism of fatty acids.<a href="#cite_note-231">[231]</a> Moreover, rs174548 in the FADS1 gene shows increased correlation with DNA methylation in people with high abundance of CD8+ T cells.<a href="#cite_note-Hawe_2022-223">[223]</a> SNP rs174548 is strongly associated with concentrations of arachidonic acid and other metabolites in fatty acid metabolism,<a href="#cite_note-232">[232]</a><a href="#cite_note-pmid24816252-233">[233]</a> blood eosinophil counts.<a href="#cite_note-234">[234]</a> and inflammatory diseases such as asthma.<a href="#cite_note-235">[235]</a> Interaction results indicated a correlation between rs174548 and asthma, providing new insights about fatty acid metabolism in CD8+ T cells with immune phenotypes.<a href="#cite_note-Hawe_2022-223">[223]</a> <div class="mw-heading mw-heading2"><h2 id="Pseudoscience">Pseudoscience</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=49" title="Edit section: Pseudoscience">edit</a>]</div> As epigenetics is in the early stages of development as a science and is surrounded by <a href="/wiki/Sensationalism" title="Sensationalism">sensationalism</a> in the public media, <a href="/wiki/David_Gorski" title="David Gorski">David Gorski</a> and geneticist <a href="/wiki/Adam_Rutherford" title="Adam Rutherford">Adam Rutherford</a> have advised caution against the proliferation of false and <a href="/wiki/Pseudoscience" title="Pseudoscience">pseudoscientific</a> conclusions by <a href="/wiki/New_age" class="mw-redirect" title="New age">new age</a> authors making unfounded suggestions that a person's genes and health can be manipulated by <a href="/wiki/Brainwashing" title="Brainwashing">mind control</a>. Misuse of the scientific term by <a href="/wiki/Quackery" title="Quackery">quack authors</a> has produced misinformation among the general public.<a href="#cite_note-science-2">[2]</a><a href="#cite_note-236">[236]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=50" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1259569809">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{clear:left;float:left;margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><a href="/wiki/File:Issoria_lathonia.jpg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/32px-Issoria_lathonia.jpg" decoding="async" width="32" height="23" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/48px-Issoria_lathonia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/64px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a><a href="/wiki/Portal:Biology" title="Portal:Biology">Biology portal</a></li><li class="portalbox-entry"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/12px-WHO_Rod.svg.png" decoding="async" width="12" height="28" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/18px-WHO_Rod.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/24px-WHO_Rod.svg.png 2x" data-file-width="107" data-file-height="250" /><a href="/wiki/Portal:Medicine" title="Portal:Medicine">Medicine portal</a></li></ul> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col"> <ul><li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a></li> <li><a href="/wiki/Biological_effects_of_radiation_on_the_epigenome" title="Biological effects of radiation on the epigenome">Biological effects of radiation on the epigenome</a></li> <li><a href="/wiki/Computational_epigenetics" title="Computational epigenetics">Computational epigenetics</a></li> <li><a href="/wiki/Contribution_of_epigenetic_modifications_to_evolution" title="Contribution of epigenetic modifications to evolution">Contribution of epigenetic modifications to evolution</a></li> <li><a href="/wiki/DAnCER_(database)" title="DAnCER (database)">DAnCER</a> database (2010)</li> <li><a href="/wiki/Epigenesis_(biology)" title="Epigenesis (biology)">Epigenesis (biology)</a></li> <li><a href="/wiki/Epigenetics_in_forensic_science" title="Epigenetics in forensic science">Epigenetics in forensic science</a></li> <li><a href="/wiki/Epigenetics_of_autoimmune_disorders" title="Epigenetics of autoimmune disorders">Epigenetics of autoimmune disorders</a></li> <li><a href="/wiki/Epiphenotyping" title="Epiphenotyping">Epiphenotyping</a></li> <li><a href="/wiki/Epigenetic_therapy" title="Epigenetic therapy">Epigenetic therapy</a></li> <li><a href="/wiki/Epigenetics_of_neurodegenerative_diseases" title="Epigenetics of neurodegenerative diseases">Epigenetics of neurodegenerative diseases</a></li> <li><a href="/wiki/Genetics" title="Genetics">Genetics</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Nutriepigenomics" title="Nutriepigenomics">Nutriepigenomics</a></li> <li><a href="/wiki/Position-effect_variegation" title="Position-effect variegation">Position-effect variegation</a></li> <li><a href="/wiki/Preformationism" title="Preformationism">Preformationism</a></li> <li><a href="/wiki/Somatic_epitype" title="Somatic epitype">Somatic epitype</a></li> <li><a href="/wiki/Synthetic_genetic_array" title="Synthetic genetic array">Synthetic genetic array</a></li> <li><a href="/wiki/Sleep_epigenetics" title="Sleep epigenetics">Sleep epigenetics</a></li> <li><a href="/wiki/Transcriptional_memory" title="Transcriptional memory">Transcriptional memory</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=51" 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 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Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-992235-2" title="Special:BookSources/978-0-19-992235-2"><bdi>978-0-19-992235-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Developing+Genome%3A+An+Introduction+to+Behavioral+Epigenetics&rft.pub=Oxford+University+Press&rft.date=2015&rft.isbn=978-0-19-992235-2&rft.aulast=Moore&rft.aufirst=DS&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988">[<a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">page needed</a>] </li> <li id="cite_note-pmid19339683-12">^ <a href="#cite_ref-pmid19339683_12-0">a</a> <a href="#cite_ref-pmid19339683_12-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBergerKouzaridesShiekhattarShilatifard2009" class="citation journal cs1">Berger SL, Kouzarides T, Shiekhattar R, Shilatifard A (April 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3959995">"An operational definition of epigenetics"</a>. Genes & Development. 23 (7): 781–3. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgad.1787609">10.1101/gad.1787609</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3959995">3959995</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19339683">19339683</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genes+%26+Development&rft.atitle=An+operational+definition+of+epigenetics&rft.volume=23&rft.issue=7&rft.pages=781-3&rft.date=2009-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3959995%23id-name%3DPMC&rft_id=info%3Apmid%2F19339683&rft_id=info%3Adoi%2F10.1101%2Fgad.1787609&rft.aulast=Berger&rft.aufirst=SL&rft.au=Kouzarides%2C+T&rft.au=Shiekhattar%2C+R&rft.au=Shilatifard%2C+A&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3959995&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> <li id="cite_note-NIH-13">^ <a href="#cite_ref-NIH_13-0">a</a> <a href="#cite_ref-NIH_13-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20191121014029/http://www.roadmapepigenomics.org/overview">"Overview"</a>. NIH Roadmap Epigenomics Project. Archived from <a rel="nofollow" class="external text" href="http://www.roadmapepigenomics.org/overview">the original</a> on 21 November 2019. Retrieved 7 December 2013.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=NIH+Roadmap+Epigenomics+Project&rft.atitle=Overview&rft_id=http%3A%2F%2Fwww.roadmapepigenomics.org%2Foverview&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> <li id="cite_note-14"><a href="#cite_ref-14">^</a> Morange M. La tentative de Nikolai Koltzoff (Koltsov) de lier génétique, embryologie et chimie physique, J. Biosciences. 2011. V. 36. P. 211-214 </li> <li id="cite_note-waddington-15"><a href="#cite_ref-waddington_15-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWaddington1942" class="citation journal cs1">Waddington CH (1942). "The epigenotype". Endeavour. 1: 18–20.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Endeavour&rft.atitle=The+epigenotype&rft.volume=1&rft.pages=18-20&rft.date=1942&rft.aulast=Waddington&rft.aufirst=CH&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> "For the purpose of a study of inheritance, the relation between phenotypes and genotypes [...] is, from a wider biological point of view, of crucial importance, since it is the kernel of the whole problem of development." </li> <li id="cite_note-16"><a href="#cite_ref-16">^</a> See <a href="/wiki/Preformationism" title="Preformationism">preformationism</a> for historical background. <a href="/wiki/Oxford_English_Dictionary" title="Oxford English Dictionary">Oxford English Dictionary</a>: "the theory that the germ is brought into existence (by successive accretions), and not merely developed, in the process of reproduction. 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Nature Neuroscience. 17 (1): 89–96. <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%2Fnn.3594">10.1038/nn.3594</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3923835">3923835</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24292232">24292232</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Neuroscience&rft.atitle=Parental+olfactory+experience+influences+behavior+and+neural+structure+in+subsequent+generations&rft.volume=17&rft.issue=1&rft.pages=89-96&rft.date=2014-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3923835%23id-name%3DPMC&rft_id=info%3Apmid%2F24292232&rft_id=info%3Adoi%2F10.1038%2Fnn.3594&rft.aulast=Dias&rft.aufirst=BG&rft.au=Ressler%2C+KJ&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3923835&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> (see comment by Gonzalo Otazu) </li> <li id="cite_note-167"><a href="#cite_ref-167">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.the-scientist.com/?articles.view/articleNo/41239/title/Epigenetics-Paper-Raises-Questions/">"Epigenetics Paper Raises Questions"</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Epigenetics+Paper+Raises+Questions&rft_id=http%3A%2F%2Fwww.the-scientist.com%2F%3Farticles.view%2FarticleNo%2F41239%2Ftitle%2FEpigenetics-Paper-Raises-Questions%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> <li id="cite_note-isbn0-19-854968-7-168"><a href="#cite_ref-isbn0-19-854968-7_168-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHoekstra2000" class="citation book cs1">Hoekstra RF (2000). 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Archived from <a rel="nofollow" class="external text" href="http://www.dundee.ac.uk/externalrelations/events/lectures.html">the original</a> on 23 May 2007.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Lecture&rft_id=http%3A%2F%2Fwww.dundee.ac.uk%2Fexternalrelations%2Fevents%2Flectures.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> <li id="cite_note-203"><a href="#cite_ref-203">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.pbs.org/wgbh/nova/transcripts/3413_genes.html">"NOVA | Transcripts | Ghost in Your Genes"</a>. PBS. 16 October 2007. Retrieved 26 July 2012.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=NOVA+%26%23124%3B+Transcripts+%26%23124%3B+Ghost+in+Your+Genes&rft.pub=PBS&rft.date=2007-10-16&rft_id=https%3A%2F%2Fwww.pbs.org%2Fwgbh%2Fnova%2Ftranscripts%2F3413_genes.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> <li id="cite_note-204"><a href="#cite_ref-204">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAndersonFeyeSchmidt-McCormackMalovic2016" class="citation journal cs1">Anderson SJ, Feye KM, Schmidt-McCormack GR, Malovic E, Mlynarczyk GS, Izbicki P, et al. (May 2016). "Off-Target drug effects resulting in altered gene expression events with epigenetic and "Quasi-Epigenetic" origins". 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Science-Based Medicine.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Science-Based+Medicine&rft.atitle=Epigenetics%3A+It+doesn%27t+mean+what+quacks+think+it+means&rft.date=2013-02-04&rft.aulast=Gorski&rft.aufirst=D&rft_id=https%3A%2F%2Fwww.sciencebasedmedicine.org%2Fepigenetics-it-doesnt-mean-what-quacks-think-it-means%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=52" title="Edit section: Further reading">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHaqueGottesmanWong2009" class="citation journal cs1">Haque FN, Gottesman II, Wong AH (May 2009). "Not really identical: epigenetic differences in monozygotic twins and implications for twin studies in psychiatry". American Journal of Medical Genetics. Part C, Seminars in Medical Genetics. 151C (2): 136–41. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fajmg.c.30206">10.1002/ajmg.c.30206</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19378334">19378334</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:205327825">205327825</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Medical+Genetics.+Part+C%2C+Seminars+in+Medical+Genetics&rft.atitle=Not+really+identical%3A+epigenetic+differences+in+monozygotic+twins+and+implications+for+twin+studies+in+psychiatry&rft.volume=151C&rft.issue=2&rft.pages=136-41&rft.date=2009-05&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A205327825%23id-name%3DS2CID&rft_id=info%3Apmid%2F19378334&rft_id=info%3Adoi%2F10.1002%2Fajmg.c.30206&rft.aulast=Haque&rft.aufirst=FN&rft.au=Gottesman%2C+II&rft.au=Wong%2C+AH&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.cdc.gov/genomics/disease/epigenetics.htm">"What is Epigenetics?"</a>. Centers for Disease Control and Prevention. 15 August 2022. Retrieved 11 September 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Centers+for+Disease+Control+and+Prevention&rft.atitle=What+is+Epigenetics%3F&rft.date=2022-08-15&rft_id=https%3A%2F%2Fwww.cdc.gov%2Fgenomics%2Fdisease%2Fepigenetics.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Epigenetics&action=edit&section=53" title="Edit section: External links">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/40px-Wiktionary-logo-en-v2.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/60px-Wiktionary-logo-en-v2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></div> <div class="side-box-text plainlist">Look up <a href="https://en.wiktionary.org/wiki/Special:Search/epigenetics" class="extiw" title="wiktionary:Special:Search/epigenetics">epigenetics</a> in Wiktionary, the free dictionary.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <a href="https://commons.wikimedia.org/wiki/Category:Epigenetics" class="extiw" title="commons:Category:Epigenetics">Epigenetics</a>.</div></div> </div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://learn.genetics.utah.edu/content/epigenetics/inheritance/">"Epigenetics & Inheritance"</a>. learn.genetics.utah.edu. Retrieved 17 April 2019.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=learn.genetics.utah.edu&rft.atitle=Epigenetics+%26+Inheritance&rft_id=https%3A%2F%2Flearn.genetics.utah.edu%2Fcontent%2Fepigenetics%2Finheritance%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEpigenetics" class="Z3988"></li> <li><a rel="nofollow" class="external text" href="http://www.epigenome.org/">The Human Epigenome Project (HEP)</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100503172418/http://www.epigenome-noe.net/index.php">The Epigenome Network of Excellence (NoE)</a></li> <li><a rel="nofollow" class="external text" href="http://www.epigenomes.ca/">Canadian Epigenetics, Environment and Health Research Consortium (CEEHRC)</a></li> <li><a rel="nofollow" class="external text" href="http://www.epigenome.eu/">The Epigenome Network of Excellence (NoE) – public international site</a></li> <li><a rel="nofollow" class="external text" href="http://discovermagazine.com/2006/nov/cover">"DNA Is Not Destiny"</a> – Discover magazine cover story</li> <li><a rel="nofollow" class="external text" href="https://www.bbc.co.uk/sn/tvradio/programmes/horizon/ghostgenes.shtml">"The Ghost In Your Genes"</a>, Horizon (2005), BBC</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080509124439/http://www.hopkinsmedicine.org/press/2002/November/epigenetics.htm">Epigenetics article</a> at Hopkins Medicine</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20110721032750/http://genome.wellcome.ac.uk/doc_WTX036556.html">Towards a global map of epigenetic variation </a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output 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href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</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/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a> of</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/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a></li> <li><a href="/wiki/Genetic_history_of_indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of indigenous peoples of the Americas">the Americas</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a></li></ul> </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/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a class="mw-selflink selflink">Epigenetics</a></li> <li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Genetic diversity</a></li> <li><a href="/wiki/Genetic_monitoring" title="Genetic monitoring">Genetic monitoring</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/He_Jiankui_affair" title="He Jiankui affair">He Jiankui affair</a></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Plant_genetics" title="Plant genetics">Plant genetics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></li> <li><a href="/wiki/List_of_genetics_research_organizations" title="List of genetics research organizations">List of genetics research organizations</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><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:Genetics" title="Category:Genetics">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="Gene_expression" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction to genetics</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/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a href="/wiki/DNA" title="DNA">DNA</a> → <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a> → <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</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:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA / hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</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/Translation_(biology)" title="Translation (biology)">Translation</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:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</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/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</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="The_development_of_phenotype" 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:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</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/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</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/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</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%">Systems</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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</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/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a href="/wiki/Morphogen" title="Morphogen">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</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/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Molecular_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Molecular_biology" title="Template:Molecular biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Molecular_biology" title="Template talk:Molecular biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Molecular_biology" title="Special:EditPage/Template:Molecular biology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Molecular_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_molecular_biology" title="History of molecular biology">History</a></li> <li><a href="/wiki/Index_of_molecular_biology_articles" title="Index of molecular biology articles">Index</a></li> <li><a href="/wiki/Glossary_of_genetics" class="mw-redirect" title="Glossary of genetics">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Overview</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/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</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/DNA_replication" title="DNA replication">DNA replication</a> (<a href="/wiki/DNA" title="DNA">DNA</a>)</li> <li><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</a> (<a href="/wiki/RNA" title="RNA">RNA</a>)</li> <li><a href="/wiki/Translation_(biology)" title="Translation (biology)">Translation</a> (<a href="/wiki/Protein" title="Protein">protein</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Element</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li> <ul><li>Genetic</li> <li>Heredity</li></ul></li></ul> <ul><li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> <ul><li><a href="/wiki/Pribnow_box" title="Pribnow box">Pribnow box</a></li> <li><a href="/wiki/TATA_box" title="TATA box">TATA box</a></li></ul></li> <li><a href="/wiki/Operon" title="Operon">Operon</a> <ul><li><a href="/wiki/Gal_operon" title="Gal operon">gal operon</a></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Trp_operon" title="Trp operon">trp operon</a></li></ul></li> <li><a href="/wiki/Intron" title="Intron">Intron</a></li> <li><a href="/wiki/Exon" title="Exon">Exon</a></li> <li><a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">Terminator</a></li> <li><a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a></li> <li><a href="/wiki/Repressor" title="Repressor">Repressor</a> <ul><li><a href="/wiki/Lac_repressor" title="Lac repressor">lac repressor</a></li> <li><a href="/wiki/Tryptophan_repressor" title="Tryptophan repressor">trp repressor</a></li></ul></li> <li><a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">Silencer</a></li> <li><a href="/wiki/Histone_methylation" title="Histone methylation">Histone methylation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Linked life</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_biology" title="Cell biology">Cell biology</a></li> <li><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a></li> <li><a href="/wiki/Computational_biology" title="Computational biology">Computational biology</a></li> <li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Medicine" title="Medicine">Functional biology/medicine</a></li> <li><a href="/wiki/Genetics" title="Genetics">Genetics</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Engineering</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%">Concepts</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/Cultured_meat" title="Cultured meat">Cultured meat</a></li> <li><a href="/wiki/Mitosis" title="Mitosis">Mitosis</a></li> <li><a href="/wiki/Cell_signaling" title="Cell signaling">Cell signalling</a></li> <li><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcriptional modification</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</a></li> <li><a href="/wiki/Dry_lab" title="Dry lab">Dry lab</a> / <a href="/wiki/Wet_lab" title="Wet lab">Wet lab</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Techniques</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_culture" title="Cell culture">Cell culture</a></li> <li><a href="/wiki/Model_organism" title="Model organism">Model organisms</a> (such as <a href="/wiki/C57BL/6" title="C57BL/6">C57BL/6 mice</a>)</li> <li>Methods <ul><li><a href="/wiki/Nucleic_acid_methods" title="Nucleic acid methods">Nucleic acid</a></li> <li><a href="/wiki/Protein_methods" title="Protein methods">Protein</a></li></ul></li> <li><a href="/wiki/Fluorescence_in_the_life_sciences" title="Fluorescence in the life sciences">Fluorescence</a>, <a href="/wiki/Pigment" title="Pigment">Pigment</a> & <a href="/wiki/Radioactivity_in_the_life_sciences" title="Radioactivity in the life sciences">Radioactivity</a></li></ul> <dl><dt>High-throughput technique ("<a href="/wiki/Omics" title="Omics">-omics</a>")</dt> <dd><a href="/wiki/DNA_microarray" title="DNA microarray">DNA microarray</a></dd> <dd><a href="/wiki/Mass_spectrometry" title="Mass spectrometry">Mass spectrometry</a></dd> <dd><a href="/wiki/Lab-on-a-chip" title="Lab-on-a-chip">Lab-on-a-chip</a></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Gene regulation</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 class="mw-selflink selflink">Epigenetic</a></li> <li><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Genetic</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational regulation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><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:Molecular_biology" title="Category:Molecular biology">Molecular biology</a></li> <li><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/16px-People_icon.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/24px-People_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/32px-People_icon.svg.png 2x" data-file-width="100" data-file-height="100" /> <a href="/wiki/Wikipedia:WikiProject_Molecular_Biology/Molecular_and_Cell_Biology" title="Wikipedia:WikiProject Molecular Biology/Molecular and Cell Biology">WikiProject</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="Branches_of_biology" 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:Branches_of_biology" title="Template:Branches of biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Branches_of_biology" title="Template talk:Branches of biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Branches_of_biology" title="Special:EditPage/Template:Branches of biology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Branches_of_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Outline_of_biology#Branches" title="Outline of biology">Branches of biology</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Aerobiology" title="Aerobiology">Aerobiology</a></li> <li><a href="/wiki/Agronomy" title="Agronomy">Agronomy</a></li> <li><a href="/wiki/Agrostology" title="Agrostology">Agrostology</a></li> <li><a href="/wiki/Anatomy" title="Anatomy">Anatomy</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Bacteriology" title="Bacteriology">Bacteriology</a></li> <li><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Biogeology" title="Biogeology">Biogeology</a></li> <li><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></li> <li><a href="/wiki/Biological_engineering" title="Biological engineering">Biological engineering</a></li> <li><a href="/wiki/Biomechanics" title="Biomechanics">Biomechanics</a></li> <li><a href="/wiki/Biophysics" title="Biophysics">Biophysics</a></li> <li><a href="/wiki/Biosemiotics" title="Biosemiotics">Biosemiotics</a></li> <li><a href="/wiki/Biostatistics" title="Biostatistics">Biostatistics</a></li> <li><a href="/wiki/Biotechnology" title="Biotechnology">Biotechnology</a></li> <li><a href="/wiki/Botany" title="Botany">Botany</a></li> <li><a href="/wiki/Cell_biology" title="Cell biology">Cell biology</a></li> <li><a href="/wiki/Cellular_microbiology" title="Cellular microbiology">Cellular microbiology</a></li> <li><a href="/wiki/Chemical_biology" title="Chemical biology">Chemical biology</a></li> <li><a href="/wiki/Chronobiology" title="Chronobiology">Chronobiology</a></li> <li><a href="/wiki/Cognitive_biology" title="Cognitive biology">Cognitive biology</a></li> <li><a href="/wiki/Computational_biology" title="Computational biology">Computational biology</a></li> <li><a href="/wiki/Conservation_biology" title="Conservation biology">Conservation biology</a></li> <li><a href="/wiki/Cryobiology" title="Cryobiology">Cryobiology</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Dendrology" title="Dendrology">Dendrology</a></li> <li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Ecology" title="Ecology">Ecology</a></li> <li><a href="/wiki/Embryology" title="Embryology">Embryology</a></li> <li><a href="/wiki/Epidemiology" title="Epidemiology">Epidemiology</a></li> <li><a class="mw-selflink selflink">Epigenetics</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Freshwater_biology" title="Freshwater biology">Freshwater biology</a></li> <li><a href="/wiki/Generative_biology" class="mw-redirect" title="Generative biology">Generative biology</a></li> <li><a href="/wiki/Genetics" title="Genetics">Genetics</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Geobiology" title="Geobiology">Geobiology</a></li> <li><a href="/wiki/Gerontology" title="Gerontology">Gerontology</a></li> <li><a href="/wiki/Herpetology" title="Herpetology">Herpetology</a></li> <li><a href="/wiki/Histology" title="Histology">Histology</a></li> <li><a href="/wiki/Human_biology" title="Human biology">Human biology</a></li> <li><a href="/wiki/Ichthyology" title="Ichthyology">Ichthyology</a></li> <li><a href="/wiki/Immunology" title="Immunology">Immunology</a></li> <li><a href="/wiki/Lipidology" title="Lipidology">Lipidology</a></li> <li><a href="/wiki/Mammalogy" title="Mammalogy">Mammalogy</a></li> <li><a href="/wiki/Marine_biology" title="Marine biology">Marine biology</a></li> <li><a href="/wiki/Mathematical_and_theoretical_biology" title="Mathematical and theoretical biology">Mathematical biology</a></li> <li><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a></li> <li><a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a></li> <li><a href="/wiki/Mycology" title="Mycology">Mycology</a></li> <li><a href="/wiki/Neontology" title="Neontology">Neontology</a></li> <li><a href="/wiki/Neuroscience" title="Neuroscience">Neuroscience</a></li> <li><a href="/wiki/Nutritional_science" title="Nutritional science">Nutrition</a></li> <li><a href="/wiki/Ornithology" title="Ornithology">Ornithology</a></li> <li><a href="/wiki/Osteology" title="Osteology">Osteology</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Parasitology" title="Parasitology">Parasitology</a></li> <li><a href="/wiki/Pathology" title="Pathology">Pathology</a></li> <li><a href="/wiki/Pharmacology" title="Pharmacology">Pharmacology</a></li> <li><a href="/wiki/Photobiology" title="Photobiology">Photobiology</a></li> <li><a href="/wiki/Phycology" title="Phycology">Phycology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Physiology" title="Physiology">Physiology</a></li> <li><a href="/wiki/Pomology" title="Pomology">Pomology</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Proteomics" title="Proteomics">Proteomics</a></li> <li><a href="/wiki/Protistology" title="Protistology">Protistology</a></li> <li><a href="/wiki/Quantum_biology" title="Quantum biology">Quantum biology</a></li> <li><a href="/wiki/Relational_biology" class="mw-redirect" title="Relational biology">Relational biology</a></li> <li><a href="/wiki/Reproductive_biology" title="Reproductive biology">Reproductive biology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Structural_biology" title="Structural biology">Structural biology</a></li> <li><a href="/wiki/Synthetic_biology" title="Synthetic biology">Synthetic biology</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Systems_biology" title="Systems biology">Systems biology</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Taxonomy</a></li> <li><a href="/wiki/Teratology" title="Teratology">Teratology</a></li> <li><a href="/wiki/Toxicology" title="Toxicology">Toxicology</a></li> <li><a href="/wiki/Virology" title="Virology">Virology</a></li> <li><a href="/wiki/Virophysics" title="Virophysics">Virophysics</a></li> <li><a href="/wiki/Xenobiology" title="Xenobiology">Xenobiology</a></li> <li><a href="/wiki/Zoology" title="Zoology">Zoology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</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/History_of_biology" title="History of biology">History of biology</a></li> <li><a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a></li> <li><a href="/wiki/Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">Timeline of biology and organic chemistry</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:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox 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