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class="vector-toc-list"> </ul> </li> <li id="toc-Immunity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Immunity"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Immunity</span> </div> </a> <ul id="toc-Immunity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gene_regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Gene_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Gene regulation</span> </div> </a> <ul id="toc-Gene_regulation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Role_in_medicine" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_medicine"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Role in medicine</span> </div> </a> <button aria-controls="toc-Role_in_medicine-sublist" class="cdx-button cdx-button--weight-quiet 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cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Human endogenous retroviruses subsection</span> </button> <ul id="toc-Human_endogenous_retroviruses-sublist" class="vector-toc-list"> <li id="toc-Classification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Classification</span> </div> </a> <ul id="toc-Classification-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Origin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Origin"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Origin</span> </div> </a> <ul id="toc-Origin-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Expression_of_HERV_proteins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Expression_of_HERV_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Expression of HERV proteins</span> </div> </a> <ul id="toc-Expression_of_HERV_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Functional_impact" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Functional_impact"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Functional impact</span> </div> </a> <ul id="toc-Functional_impact-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Activation_by_exogenous_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Activation_by_exogenous_viruses"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.5</span> <span>Activation by exogenous viruses</span> </div> </a> <ul id="toc-Activation_by_exogenous_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Immune_response_to_HERVs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Immune_response_to_HERVs"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.6</span> <span>Immune response to HERVs</span> </div> </a> <ul id="toc-Immune_response_to_HERVs-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Techniques_for_characterizing_ERVs" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Techniques_for_characterizing_ERVs"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Techniques for characterizing ERVs</span> </div> </a> <button aria-controls="toc-Techniques_for_characterizing_ERVs-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Techniques for characterizing ERVs subsection</span> </button> <ul id="toc-Techniques_for_characterizing_ERVs-sublist" class="vector-toc-list"> <li id="toc-Whole_genome_sequencing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Whole_genome_sequencing"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Whole genome sequencing</span> </div> </a> <ul id="toc-Whole_genome_sequencing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Chromatin_immunoprecipitation_with_sequencing_(ChIP-seq)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Chromatin_immunoprecipitation_with_sequencing_(ChIP-seq)"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Chromatin immunoprecipitation with sequencing (ChIP-seq)</span> </div> </a> <ul id="toc-Chromatin_immunoprecipitation_with_sequencing_(ChIP-seq)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Applications" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Applications"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Applications</span> </div> </a> <button aria-controls="toc-Applications-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Applications subsection</span> </button> <ul id="toc-Applications-sublist" class="vector-toc-list"> <li id="toc-Constructing_phylogenies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Constructing_phylogenies"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Constructing phylogenies</span> </div> </a> <ul id="toc-Constructing_phylogenies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Designating_the_age_of_provirus_and_the_time_points_of_species_separation_events" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Designating_the_age_of_provirus_and_the_time_points_of_species_separation_events"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Designating the age of provirus and the time points of species separation events</span> </div> </a> <ul id="toc-Designating_the_age_of_provirus_and_the_time_points_of_species_separation_events-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Further_research" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_research"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Further research</span> </div> </a> <button aria-controls="toc-Further_research-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Further research subsection</span> </button> <ul id="toc-Further_research-sublist" class="vector-toc-list"> <li id="toc-Epigenetic_variability" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Epigenetic_variability"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Epigenetic variability</span> </div> </a> <ul id="toc-Epigenetic_variability-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Immunological_problems_of_xenotransplantation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Immunological_problems_of_xenotransplantation"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Immunological problems of xenotransplantation</span> </div> </a> <ul id="toc-Immunological_problems_of_xenotransplantation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Risk_factors_of_HERVs_in_gene_therapy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Risk_factors_of_HERVs_in_gene_therapy"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.3</span> <span>Risk factors of HERVs in gene therapy</span> </div> </a> <ul id="toc-Risk_factors_of_HERVs_in_gene_therapy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-HERV_gene_expression" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#HERV_gene_expression"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.4</span> <span>HERV gene expression</span> </div> </a> <ul id="toc-HERV_gene_expression-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > 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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 17 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-17" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">17 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3_%D8%B1%D8%A7%D8%AC%D8%B9_%D8%AF%D8%A7%D8%AE%D9%84%D9%8A" title="فيروس راجع داخلي – Arabic" lang="ar" hreflang="ar" data-title="فيروس راجع داخلي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Retrovirus_endogen" title="Retrovirus endogen – Catalan" lang="ca" hreflang="ca" data-title="Retrovirus endogen" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Endogen_retrovirus" title="Endogen retrovirus – Danish" lang="da" hreflang="da" data-title="Endogen retrovirus" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Endogenes_Retrovirus" title="Endogenes Retrovirus – German" lang="de" hreflang="de" data-title="Endogenes Retrovirus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Endogeensed_retroviirused" title="Endogeensed retroviirused – Estonian" lang="et" hreflang="et" data-title="Endogeensed retroviirused" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Retrovirus_end%C3%B3geno" title="Retrovirus endógeno – Spanish" lang="es" hreflang="es" data-title="Retrovirus endógeno" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/R%C3%A9trovirus_endog%C3%A8ne" title="Rétrovirus endogène – French" lang="fr" hreflang="fr" data-title="Rétrovirus endogène" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Retrovirus_end%C3%B3xeno" title="Retrovirus endóxeno – Galician" lang="gl" hreflang="gl" data-title="Retrovirus endóxeno" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Endogenes_retrovirus" title="Endogenes retrovirus – Latin" lang="la" hreflang="la" data-title="Endogenes retrovirus" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lmo mw-list-item"><a href="https://lmo.wikipedia.org/wiki/Retrovirus_endogen" title="Retrovirus endogen – Lombard" lang="lmo" hreflang="lmo" data-title="Retrovirus endogen" data-language-autonym="Lombard" data-language-local-name="Lombard" class="interlanguage-link-target"><span>Lombard</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%86%85%E5%9C%A8%E6%80%A7%E3%83%AC%E3%83%88%E3%83%AD%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="内在性レトロウイルス – Japanese" lang="ja" hreflang="ja" data-title="内在性レトロウイルス" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Retrowirusy_endogenne" title="Retrowirusy endogenne – Polish" lang="pl" hreflang="pl" data-title="Retrowirusy endogenne" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Retrov%C3%ADrus_end%C3%B3geno" title="Retrovírus endógeno – Portuguese" lang="pt" hreflang="pt" data-title="Retrovírus endógeno" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Endogeeninen_retrovirus" title="Endogeeninen retrovirus – Finnish" lang="fi" hreflang="fi" data-title="Endogeeninen retrovirus" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Endogent_retrovirus" title="Endogent retrovirus – Swedish" lang="sv" hreflang="sv" data-title="Endogent retrovirus" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%95%D0%BD%D0%B4%D0%BE%D0%B3%D0%B5%D0%BD%D0%BD%D1%96_%D1%80%D0%B5%D1%82%D1%80%D0%BE%D0%B2%D1%96%D1%80%D1%83%D1%81%D0%B8" title="Ендогенні ретровіруси – Ukrainian" lang="uk" hreflang="uk" data-title="Ендогенні ретровіруси" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%85%A7%E6%BA%90%E6%80%A7%E5%8F%8D%E8%BD%89%E9%8C%84%E7%97%85%E6%AF%92" title="內源性反轉錄病毒 – Chinese" lang="zh" hreflang="zh" data-title="內源性反轉錄病毒" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a 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searchaux" style="display:none">Inherited retrovirus encoded in an organism's genome</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">"HERV" redirects here. For other uses, see <a href="/wiki/Herv_(disambiguation)" class="mw-redirect mw-disambig" title="Herv (disambiguation)">Herv (disambiguation)</a>.</div><style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Cleanup plainlinks metadata ambox ambox-style ambox-Cleanup" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article may <b>require <a href="/wiki/Wikipedia:Cleanup" title="Wikipedia:Cleanup">cleanup</a></b> to meet Wikipedia's <a href="/wiki/Wikipedia:Manual_of_Style" title="Wikipedia:Manual of Style">quality standards</a>. The specific problem is: <b>Some links missing, also check for overlink.</b><span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Endogenous_retrovirus" title="Special:EditPage/Endogenous retrovirus">improve this article</a> if you can.</span> <span class="date-container"><i>(<span class="date">February 2023</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Classes_of_ERVs.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/31/Classes_of_ERVs.svg/260px-Classes_of_ERVs.svg.png" decoding="async" width="260" height="299" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/31/Classes_of_ERVs.svg/390px-Classes_of_ERVs.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/31/Classes_of_ERVs.svg/520px-Classes_of_ERVs.svg.png 2x" data-file-width="630" data-file-height="725" /></a><figcaption><a href="/wiki/Dendrogram" title="Dendrogram">Dendrogram</a> of various classes of <a href="/wiki/Endogeny_(biology)" title="Endogeny (biology)">endogenous</a> retroviruses</figcaption></figure> <p><b>Endogenous retroviruses</b> (<b>ERVs</b>) are <a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">endogenous viral elements</a> in the <a href="/wiki/Genome" title="Genome">genome</a> that closely resemble and can be derived from <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>. They are abundant in the genomes of <a href="/wiki/Gnathostomata" title="Gnathostomata">jawed vertebrates</a>, and they comprise up to 5–8% of the human genome (lower estimates of ~1%).<sup id="cite_ref-Belshaw2004_1-0" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>An ERV is a vertically inherited <a href="/wiki/Provirus" title="Provirus">proviral</a> sequence and a subclass of a type of <a href="/wiki/Gene" title="Gene">gene</a> called a <a href="/wiki/Transposon" class="mw-redirect" title="Transposon">transposon</a>, which can normally be packaged and moved within the genome to serve a vital role in <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> and in <a href="/wiki/Gene_regulation" class="mw-redirect" title="Gene regulation">regulation</a>.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> ERVs, however, lack most transposon functions, are typically not infectious, and are often defective genomic remnants of the retroviral replication cycle.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Nomenclature_for_endogenous_retrovi_6-0" class="reference"><a href="#cite_note-Nomenclature_for_endogenous_retrovi-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> They are distinguished as <a href="/wiki/Germline" title="Germline">germline</a> <a href="/wiki/Provirus" title="Provirus">provirus</a> <a href="/wiki/Retroelements" class="mw-redirect" title="Retroelements">retroelements</a> due to their integration and <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse-transcription</a> into the nuclear genome of the host cell. </p><p>Researchers have suggested that retroviruses evolved from a type of transposon called a <a href="/wiki/Retrotransposon" title="Retrotransposon">retrotransposon</a>, a Class I element;<sup id="cite_ref-Rebollo12_7-0" class="reference"><a href="#cite_note-Rebollo12-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> these genes can mutate and instead of moving to another location in the genome, they can become exogenous or pathogenic. This means that not all ERVs may have originated as an insertion by a retrovirus but that some may have been the source for the genetic information in the retroviruses they resemble.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> When integration of viral DNA occurs in the germ-line, it can give rise to an ERV, which can later become fixed in the gene pool of the host population.<sup id="cite_ref-Belshaw2004_1-1" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Johnson_10254–10260_9-0" class="reference"><a href="#cite_note-Johnson_10254–10260-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Formation">Formation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=1" title="Edit section: Formation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The replication cycle of a <a href="/wiki/Retrovirus" title="Retrovirus">retrovirus</a> entails the insertion ("integration") of a DNA copy of the viral <a href="/wiki/Genome" title="Genome">genome</a> into the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nuclear</a> genome of the <a href="/wiki/Host_(biology)" title="Host (biology)">host</a> <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>. Most retroviruses infect <a href="/wiki/Somatic_cells" class="mw-redirect" title="Somatic cells">somatic cells</a>, but occasional infection of <a href="/wiki/Germline" title="Germline">germline</a> cells (cells that produce eggs and sperm) can also occur. Rarely, retroviral integration may occur in a germline cell that goes on to develop into a viable organism. This organism will carry the inserted retroviral genome as an integral part of its own genome—an "endogenous" retrovirus (ERV) that may be <a href="/wiki/Heredity" title="Heredity">inherited</a> by its <a href="/wiki/Offspring" title="Offspring">offspring</a> as a novel <a href="/wiki/Allele" title="Allele">allele</a>. Many ERVs have persisted in the genome of their hosts for millions of years. However, most of these have acquired inactivating <a href="/wiki/Mutation" title="Mutation">mutations</a> during host <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> and are no longer capable of producing the virus. ERVs can also be partially excised from the genome by a process known as recombinational deletion, in which <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a> between the identical sequences that flank newly integrated retroviruses results in deletion of the internal, <a href="/wiki/Protein" title="Protein">protein</a>-coding regions of the viral genome. </p><p>The general retrovirus genome consists of three genes vital for the invasion, replication, escape, and spreading of its viral genome. These three genes are gag (encodes for structural proteins for the viral core), pol (encodes for <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, <a href="/wiki/Integrase" title="Integrase">integrase</a>, and <a href="/wiki/Protease" title="Protease">protease</a>), and env (encodes for <a href="/wiki/Viral_coat_protein" class="mw-redirect" title="Viral coat protein">coat proteins</a> for the virus's exterior). These viral proteins are encoded as <a href="/wiki/Polyprotein" class="mw-redirect" title="Polyprotein">polyproteins</a>. In order to carry out their life cycle, the retrovirus relies heavily on the host cell's machinery. Protease degrades peptide bonds of the viral polyproteins, making the separate proteins functional. Reverse transcriptase functions to synthesize viral DNA from the viral RNA in the host cell's cytoplasm before it enters the nucleus. Integrase guides the integration of viral DNA into the host genome.<sup id="cite_ref-Johnson_10254–10260_9-1" class="reference"><a href="#cite_note-Johnson_10254–10260-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Over time, the genome of ERVs not only acquire point mutations, but also shuffle and recombine with other ERVs.<sup id="cite_ref-pmid26800882_11-0" class="reference"><a href="#cite_note-pmid26800882-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> ERVs with a decayed sequence for the <i>env</i> become more likely to propagate.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_genomic_evolution">Role in genomic evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=2" title="Edit section: Role in genomic evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Integration_of_viral_DNA_into_host_genome.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Integration_of_viral_DNA_into_host_genome.png/220px-Integration_of_viral_DNA_into_host_genome.png" decoding="async" width="220" height="178" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Integration_of_viral_DNA_into_host_genome.png/330px-Integration_of_viral_DNA_into_host_genome.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/49/Integration_of_viral_DNA_into_host_genome.png/440px-Integration_of_viral_DNA_into_host_genome.png 2x" data-file-width="858" data-file-height="694" /></a><figcaption>Diagram displaying the integration of viral DNA into a host genome</figcaption></figure> <p>Endogenous retroviruses can play an active role in shaping genomes. Most studies in this area have focused on the genomes of humans and higher primates, but other vertebrates, such as mice and sheep, have also been studied in depth.<sup id="cite_ref-Li12_13-0" class="reference"><a href="#cite_note-Li12-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-no7_14-0" class="reference"><a href="#cite_note-no7-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Black10_15-0" class="reference"><a href="#cite_note-Black10-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid15574851_16-0" class="reference"><a href="#cite_note-pmid15574851-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The long terminal repeat (<a href="/wiki/Long_terminal_repeat" title="Long terminal repeat">LTR</a>) sequences that flank ERV genomes frequently act as alternate <a href="/wiki/Gene_promoter" class="mw-redirect" title="Gene promoter">promoters</a> and <a href="/wiki/Gene_enhancer" class="mw-redirect" title="Gene enhancer">enhancers</a>, often contributing to the <a href="/wiki/Alternative_splicing" title="Alternative splicing">transcriptome</a> by producing tissue-specific variants. In addition, the retroviral <a href="/wiki/Protein" title="Protein">proteins</a> themselves have been co-opted to serve novel host functions, particularly in reproduction and development. Recombination between homologous retroviral sequences has also contributed to gene shuffling and the generation of genetic variation. Furthermore, in the instance of potentially antagonistic effects of retroviral sequences, repressor genes have co-evolved to combat them. </p><p>About 90% of endogenous retroviruses are solo LTRs, lacking all <a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frames</a> (ORFs). Solo LTRs and LTRs associated with complete retroviral sequences have been shown to act as transcriptional elements on host genes. Their range of action is mainly by insertion into the 5' <a href="/wiki/Untranslated_region" title="Untranslated region">untranslated regions (UTRs)</a> of protein coding genes; however, they have been known to act upon genes up to 70–100 <a href="/wiki/Base_pair#Length_measurements" title="Base pair">kb</a> away.<sup id="cite_ref-Li12_13-1" class="reference"><a href="#cite_note-Li12-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lagemaat03_18-0" class="reference"><a href="#cite_note-Lagemaat03-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kovalskaya06_19-0" class="reference"><a href="#cite_note-Kovalskaya06-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> The majority of these elements are inserted in the sense direction (5' to 3') to their corresponding genes, but there has been evidence<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> of LTRs acting in the antisense direction (3' to 5') and as a bidirectional promoter for neighboring genes.<sup id="cite_ref-no11_21-0" class="reference"><a href="#cite_note-no11-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-no12_22-0" class="reference"><a href="#cite_note-no12-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> In a few cases, the LTR functions as the major promoter for the gene. </p><p>For example, in humans, amylase alpha 1C (AMY1C) has a complete ERV sequence in its promoter region; the associated LTR confers salivary specific expression of the digestive <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> <a href="/wiki/Amylase" title="Amylase">amylase</a>.<sup id="cite_ref-no13_23-0" class="reference"><a href="#cite_note-no13-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Also, the primary promoter for <a href="/wiki/BAAT" title="BAAT">bile acid-CoA:amino acid N-acyltransferase</a> (BAAT), which codes for an enzyme that is integral in bile metabolism, is of LTR origin.<sup id="cite_ref-Lagemaat03_18-1" class="reference"><a href="#cite_note-Lagemaat03-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cohen09_24-0" class="reference"><a href="#cite_note-Cohen09-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>The insertion of a solo ERV-9 LTR may have produced a functional open reading frame, causing the rebirth of the human <a href="/wiki/IRGM" title="IRGM">immunity related GTPase gene</a> (IRGM).<sup id="cite_ref-no15_25-0" class="reference"><a href="#cite_note-no15-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> ERV insertions have also been shown to generate alternative splice sites either by direct integration into the gene, as with the human leptin hormone receptor, or driven by the expression of an upstream LTR, as with the phospholipase A-2 like protein.<sup id="cite_ref-Jern08_26-0" class="reference"><a href="#cite_note-Jern08-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>Most of the time, however, the LTR functions as one of many alternate promoters, often conferring tissue-specific expression related to reproduction and development. In fact, 64% of known LTR-promoted transcription variants are <a href="/wiki/Gene_expression" title="Gene expression">expressed</a> in reproductive tissues.<sup id="cite_ref-no16_27-0" class="reference"><a href="#cite_note-no16-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> For example, the gene CYP19 codes for <a href="/wiki/Aromatase" title="Aromatase">aromatase</a> P450, an important enzyme for estrogen synthesis, that is normally expressed in the brain and reproductive organs of most mammals.<sup id="cite_ref-Lagemaat03_18-2" class="reference"><a href="#cite_note-Lagemaat03-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> However, in primates, an LTR-promoted transcriptional variant confers expression to the placenta and is responsible for controlling estrogen levels during pregnancy.<sup id="cite_ref-Lagemaat03_18-3" class="reference"><a href="#cite_note-Lagemaat03-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Furthermore, the <a href="/wiki/NAIP_(gene)" title="NAIP (gene)">neuronal apoptosis inhibitory protein</a> (NAIP), normally widespread, has an LTR of the human ERV-P (HERV-P) family acting as a promoter that confers expression to the testis and prostate.<sup id="cite_ref-no17_28-0" class="reference"><a href="#cite_note-no17-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Other proteins, such as nitric oxide synthase 3 (<a href="/wiki/NOS3" class="mw-redirect" title="NOS3">NOS3</a>), interleukin-2 receptor B (<a href="/wiki/IL2RB" title="IL2RB">IL-2RB</a>), and another mediator of estrogen synthesis, <a href="/wiki/HSD17B1" title="HSD17B1">HSD17B1</a>, are also alternatively regulated by LTRs that confer placental expression, but their specific functions are not yet known.<sup id="cite_ref-Cohen09_24-1" class="reference"><a href="#cite_note-Cohen09-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-no18_29-0" class="reference"><a href="#cite_note-no18-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> The high degree of reproductive expression is thought to be an after effect of the method by which they were endogenized; however, this also may be due to a lack of <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> in germ-line tissues.<sup id="cite_ref-Cohen09_24-2" class="reference"><a href="#cite_note-Cohen09-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>The best-characterized instance of placental protein expression comes not from an alternatively promoted host gene but from a complete co-option of a retroviral protein. Retroviral fusogenic env proteins, which play a role in the entry of the virion into the host cell, have had an important impact on the development of the mammalian <a href="/wiki/Placenta" title="Placenta">placenta</a>. In mammals, intact env proteins called <a href="/wiki/ERVWE1" class="mw-redirect" title="ERVWE1">syncytins</a> are responsible for the formation and function of <a href="/wiki/Syncytiotrophoblasts" class="mw-redirect" title="Syncytiotrophoblasts">syncytiotrophoblasts</a>.<sup id="cite_ref-Black10_15-1" class="reference"><a href="#cite_note-Black10-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> These multinucleated cells are mainly responsible for maintaining nutrient exchange and separating the fetus from the mother's immune system.<sup id="cite_ref-Black10_15-2" class="reference"><a href="#cite_note-Black10-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> It has been suggested that the selection and fixation of these proteins for this function have played a critical role in the evolution of <a href="/wiki/Viviparity" title="Viviparity">viviparity</a>.<sup id="cite_ref-pmid8995601_30-0" class="reference"><a href="#cite_note-pmid8995601-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>In addition, the insertion of ERVs and their respective LTRs have the potential to induce chromosomal rearrangement due to recombination between viral sequences at inter-chromosomal loci. These rearrangements have been shown to induce gene duplications and deletions that largely contribute to genome plasticity and dramatically change the dynamic of gene function.<sup id="cite_ref-no20_31-0" class="reference"><a href="#cite_note-no20-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Furthermore, retroelements in general are largely prevalent in rapidly evolving, mammal-specific gene families whose function is largely related to the response to stress and external stimuli.<sup id="cite_ref-Lagemaat03_18-4" class="reference"><a href="#cite_note-Lagemaat03-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> In particular, both human <a href="/wiki/Class_I_MHC" class="mw-redirect" title="Class I MHC">class I</a> and <a href="/wiki/Class_II_MHC" class="mw-redirect" title="Class II MHC">class II MHC</a> genes have a high density of HERV elements as compared to other multi-locus-gene families.<sup id="cite_ref-Jern08_26-1" class="reference"><a href="#cite_note-Jern08-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> It has been shown that HERVs have contributed to the formation of extensively duplicated duplicon blocks that make up the human leukocyte antigen (HLA) class 1 family of genes.<sup id="cite_ref-Dawkins99_32-0" class="reference"><a href="#cite_note-Dawkins99-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> More specifically, HERVs primarily occupy regions within and between the break points of these blocks, suggesting that considerable duplication and deletion events, typically associated with unequal crossover, facilitated their formation.<sup id="cite_ref-no22_33-0" class="reference"><a href="#cite_note-no22-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The generation of these blocks, inherited as immunohaplotypes, act as a protective polymorphism against a wide range of antigens that may have imbued humans with an advantage over other primates.<sup id="cite_ref-Dawkins99_32-1" class="reference"><a href="#cite_note-Dawkins99-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>The characteristic of <a href="/wiki/Placenta" title="Placenta">placentas</a> being very evolutionary distinct organs between different species has been suggested to result from the <a href="/wiki/Co-option" title="Co-option">co-option</a> of ERV enhancers. Regulatory mutations, instead of mutations in genes that encode for <a href="/wiki/Hormone" title="Hormone">hormones</a> and <a href="/wiki/Growth_factor" title="Growth factor">growth factors</a>, support the known evolution of placental morphology, especially since the majority of hormone and growth factor genes are expressed in response to pregnancy, not during placental development. Researchers studied the regulatory landscape of placental development between the rat and mouse, two closely related species. This was done by mapping all regulatory elements of the rat <a href="/wiki/Trophoblast" title="Trophoblast">trophoblast</a> <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a> (TSCs) and comparing them to their <a href="/wiki/Ortholog" class="mw-redirect" title="Ortholog">orthologs</a> in mouse TSCs. TSCs were observed because they reflect the initial cells that develop in the fetal placenta. Regardless of their tangible similarities, enhancer and repressed regions were mostly species-specific. However, most promoter sequences were conserved between mouse and rat. In conclusion to their study, researchers proposed that ERVs influenced species-specific placental evolution through mediation of placental growth, <a href="/wiki/Immunosuppression" title="Immunosuppression">immunosuppression</a>, and <a href="/wiki/Cell_fusion" title="Cell fusion">cell fusion</a>.<sup id="cite_ref-Chuong_325–329_34-0" class="reference"><a href="#cite_note-Chuong_325–329-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Another example of ERV exploiting cellular mechanisms is <a href="/wiki/P53" title="P53">p53</a>, a <a href="/wiki/Tumor_suppressor_gene" title="Tumor suppressor gene">tumor suppressor gene</a> (TSG). DNA damage and cellular stress induces the p53 pathway, which results in cell <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>. Using chromatin immunoprecipitation with sequencing, thirty-percent of all p53-binding sites were located within copies of a few primate-specific ERV families. A study suggested that this benefits retroviruses because p53's mechanism provides a rapid induction of transcription, which leads to the exit of viral RNA from the host cell.<sup id="cite_ref-Rebollo12_7-1" class="reference"><a href="#cite_note-Rebollo12-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Finally, the insertion of ERVs or ERV elements into genic regions of host DNA, or overexpression of their transcriptional variants, has a much higher potential to produce deleterious effects than positive ones. Their appearance into the genome has created a <a href="/wiki/Coevolution" title="Coevolution">coevolutionary</a> dynamic that proliferated the duplication and expansion of repressor genes. The most clear-cut example of this involves the rapid duplication and proliferation of tandem <a href="/wiki/Zinc-finger" class="mw-redirect" title="Zinc-finger">zinc-finger</a> genes in mammal genomes. Zinc-finger genes, particularly those that include a (Krüppel associated box) <a href="/wiki/KRAB_domain" class="mw-redirect" title="KRAB domain">KRAB domain</a>, exist in high copy number in vertebrate genomes, and their range of functions are limited to transcriptional roles.<sup id="cite_ref-no23_35-0" class="reference"><a href="#cite_note-no23-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> It has been shown in mammals, however, that the diversification of these genes was due to multiple duplication and fixation events in response to new retroviral sequences or their endogenous copies to repress their transcription.<sup id="cite_ref-Kovalskaya06_19-1" class="reference"><a href="#cite_note-Kovalskaya06-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_disease">Role in disease</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=3" title="Edit section: Role in disease"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The majority of ERVs that occur in vertebrate genomes are ancient, inactivated by mutation, and have reached genetic <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixation</a> in their host species. For these reasons, they are extremely unlikely to have negative effects on their hosts except under unusual circumstances. Nevertheless, it is clear from studies in birds and non-human mammal species including mice, cats and <a href="/wiki/Koala_retrovirus" title="Koala retrovirus">koalas</a>, that younger (i.e., more recently integrated) ERVs can be associated with disease.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The number of active ERVs in the genome of mammals is negatively related to their body size, suggesting a contribution to <a href="/wiki/Peto%27s_paradox" title="Peto's paradox">Peto's paradox</a> through cancer pathogenesis.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> This has led researchers to propose a role for ERVs in several forms of human cancer and <a href="/wiki/Autoimmune_disease" title="Autoimmune disease">autoimmune disease</a>, although conclusive evidence is lacking.<sup id="cite_ref-no1_38-0" class="reference"><a href="#cite_note-no1-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid17661701_40-0" class="reference"><a href="#cite_note-pmid17661701-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neurological_disorders">Neurological disorders</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=4" title="Edit section: Neurological disorders"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In humans, ERVs have been proposed to be involved in <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a> (MS). A specific association between MS and the <a href="/wiki/ERVWE1" class="mw-redirect" title="ERVWE1">ERVWE1</a>, or "syncytin" gene, which is derived from an ERV insertion, has been reported, along with the presence of an "MS-associated retrovirus" (MSRV), in patients with the disease.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> HERVs have also been implicated in amyotrophic lateral sclerosis (<a href="/wiki/ALS" title="ALS">ALS)</a><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> and addiction.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p>In 2004 it was reported that antibodies to HERVs were found in greater frequency in the <a href="/wiki/Serous_fluid" title="Serous fluid">sera</a> of people with <a href="/wiki/Schizophrenia" title="Schizophrenia">schizophrenia</a>. Additionally, the <a href="/wiki/Cerebrospinal_fluid" title="Cerebrospinal fluid">cerebrospinal fluid</a> of people with recent onset schizophrenia contained levels of a retroviral marker, <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, four times higher than control subjects.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Researchers continue to look at a possible link between HERVs and schizophrenia, with the additional possibility of a triggering <a href="/wiki/Pathogenic_theory_of_schizophrenia" class="mw-redirect" title="Pathogenic theory of schizophrenia">infection inducing schizophrenia</a>.<sup id="cite_ref-fox_49-0" class="reference"><a href="#cite_note-fox-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Immunity">Immunity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=5" title="Edit section: Immunity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>ERVs have been found to be associated to disease not only through disease-causing relations, but also through immunity. The frequency of ERVs in LTRs likely correlates to viral adaptations to take advantage of immunity signaling pathways that promote viral transcription and replication. A study done in 2016 investigated the benefit of ancient viral DNA integrated into a host through gene regulation networks induced by <a href="/wiki/Interferons" class="mw-redirect" title="Interferons">interferons</a>, a branch of innate immunity.<sup id="cite_ref-Patience97_50-0" class="reference"><a href="#cite_note-Patience97-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> These <a href="/wiki/Cytokines" class="mw-redirect" title="Cytokines">cytokines</a> are first to respond to viral infection and are also important in immunosurveillance for malignant cells.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> ERVs are predicted to act as cis-regulatory elements, but much of the adaptive consequences for certain physiological functions is still unknown. There is data that supports the general role of ERVs in the regulation of human interferon (IFN) response, specifically to <a href="/wiki/Interferon-gamma" class="mw-redirect" title="Interferon-gamma">interferon-gamma</a> (IFN-G). For example, IFN-stimulated genes were found to be greatly enriched with ERVs bound by <a href="/wiki/Signal_transducer_and_activator_of_transcription_1" class="mw-redirect" title="Signal transducer and activator of transcription 1">signal transducer and activator of transcription 1</a> (STAT1) and/or <a href="/wiki/Interferon_regulatory_factor" class="mw-redirect" title="Interferon regulatory factor">Interferon regulatory factor</a> (IRF1) in <a href="/wiki/CD14%2B" class="mw-redirect" title="CD14+">CD14+</a> macrophages.<sup id="cite_ref-Belshaw2004_1-2" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>HERVs also play various roles shaping the human <a href="/wiki/Innate_immunity" class="mw-redirect" title="Innate immunity">innate immunity</a> response, with some sequences activating the system and others suppressing it. They may also protect from exogenous retroviral infections: the virus-like transcripts can activate <a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern recognition receptors</a>, and the proteins can interfere with active retroviruses. A gag protein from HERV-K(HML2) is shown to mix with HIV Gag, impairing HIV capsid formation as a result.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Gene_regulation">Gene regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=6" title="Edit section: Gene regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Another idea proposed was that ERVs from the same family played a role in recruiting multiple genes into the same network of regulation. It was found that MER41 elements provided additional redundant regulatory enhancement to the genes located near STAT1 binding sites.<sup id="cite_ref-Belshaw2004_1-3" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_medicine">Role in medicine</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=7" title="Edit section: Role in medicine"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Porcine_endogenous_retrovirus">Porcine endogenous retrovirus</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=8" title="Edit section: Porcine endogenous retrovirus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Xenotransplantation#Porcine_endogenous_retroviruses" title="Xenotransplantation">Xenotransplantation § Porcine endogenous retroviruses</a></div> <p>For humans, <a href="/wiki/Porcine" class="mw-redirect" title="Porcine">porcine</a> endogenous retroviruses (PERVs) pose a concern when using porcine tissues and organs in xenotransplantation, the transplanting of living cells, tissues, and organs from an organism of one species to an organism of different species. Although pigs are generally the most suitable donors to treat human organ diseases due to practical, financial, safety, and ethical reasons,<sup id="cite_ref-Patience97_50-1" class="reference"><a href="#cite_note-Patience97-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> PERVs previously could not be removed from pigs, due to their viral ability to integrate into the host genome and passed into offspring, until the 2017, when one lab, using <a href="/wiki/CRISPR-Cas9" class="mw-redirect" title="CRISPR-Cas9">CRISPR-Cas9</a>, removed all 62 retroviruses from the pig genome.<sup id="cite_ref-pmid28798043_53-0" class="reference"><a href="#cite_note-pmid28798043-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> The consequences of cross-species transmission remain unexplored and have dangerous potential.<sup id="cite_ref-Tang16_54-0" class="reference"><a href="#cite_note-Tang16-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>Researchers have indicated that infection of human tissues by PERVs is very possible, especially in immunosuppressed individuals. An immunosuppressed condition could potentially permit a more rapid and tenacious replication of viral DNA, and would later have less difficulty adapting to human-to-human transmission. Although known infectious pathogens present in the donor organ/tissue can be eliminated by breeding pathogen-free herds, unknown retroviruses can be present in the donor. These retroviruses are often latent and asymptomatic in the donor, but can become active in the recipient. Some examples of endogenous viruses that can infect and multiply in human cells are from baboons (BaEV), cats (RD114), and mice.<sup id="cite_ref-Patience97_50-2" class="reference"><a href="#cite_note-Patience97-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>There are three different classes of PERVs, PERV-A, PERV-B, and PERV-C. PERV-A and PERV-B are <a href="/wiki/Polytropic" class="mw-redirect" title="Polytropic">polytropic</a> and can infect human cells in vitro, while PERV-C is <a href="/wiki/Ecotropic" class="mw-redirect" title="Ecotropic">ecotropic</a> and does not replicate on human cells. The major differences between the classes is in the receptor binding domain of the <i>env</i> protein and the LTRs that influence the replication of each class. PERV-A and PERV-B display LTRs that have repeats in the <a href="/wiki/Small_nucleolar_RNA_U3" title="Small nucleolar RNA U3">U3</a> region. However, PERV-A and PERV-C show repeatless LTRs. Researchers found that PERVs in culture actively adapted to the repeat structure of their LTR in order to match the best replication performance a host cell could perform. At the end of their study, researchers concluded that repeatless PERV LTR evolved from the repeat-harboring LTR. This was likely to have occurred from insertional mutation and was proven through use of data on LTR and <i>env</i>/Env. It is thought that the generation of repeatless LTRs could be reflective of an adaptation process of the virus, changing from an exogenous to an endogenous lifestyle.<sup id="cite_ref-Tönjes03_55-0" class="reference"><a href="#cite_note-Tönjes03-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>A clinical trial study performed in 1999 sampled 160 patients who were treated with different living pig tissues and observed no evidence of a persistent PERV infection in 97% of the patients from whom a sufficient amount of DNA was available to PCR for amplification of PERV sequences. This study stated found that retrospective studies are limited to finding the true incidence of infection or associated clinical symptoms. It suggested using closely monitored prospective trials which would provide a more complete and detailed evaluation of the possible cross-species PERV transmission and a comparison of the PERV.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Human_endogenous_retroviruses">Human endogenous retroviruses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=9" title="Edit section: Human endogenous retroviruses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"HERV" redirects here. For other uses, see <a href="/wiki/Herv_(disambiguation)" class="mw-redirect mw-disambig" title="Herv (disambiguation)">Herv (disambiguation)</a>.</div> <p>HERVs comprise a significant part of the <a href="/wiki/Human_genome" title="Human genome">human genome</a>, with approximately 98,000 ERV elements and fragments making up 5–8%.<sup id="cite_ref-Belshaw2004_1-4" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> According to a study published in 2005, no HERVs capable of replication had been identified; all appeared to be defective, containing major deletions or nonsense mutations (not true for HERV-K). This is because most HERVs are merely traces of original viruses, having first integrated millions of years ago. An analysis of HERV integrations is ongoing as part of the <a href="/wiki/100,000_Genomes_Project" title="100,000 Genomes Project">100,000 Genomes Project</a>.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>A 2023 study found HERV can become awakened from dormant states and contribute to <a href="/wiki/Senescence" title="Senescence">aging</a> which could <a href="/wiki/Life_extension" title="Life extension">be blocked</a> by <a href="/wiki/Neutralizing_antibody" title="Neutralizing antibody">neutralizing antibodies</a>.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p><p>HERVs were originally discovered when human genomic libraries were screened under low-stringency conditions using either probes from animal retroviruses or by using <a href="/wiki/Oligonucleotide" title="Oligonucleotide">oligonucleotides</a> with similarity to virus sequences.<sup id="cite_ref-Belshaw2004_1-5" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Classification">Classification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=10" title="Edit section: Classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>HERVs are classified based on their homologies to animal retroviruses. Families belonging to <b>Class I</b> are similar in sequence to mammalian <i><a href="/wiki/Gammaretrovirus" title="Gammaretrovirus">Gammaretroviruses</a></i> (type C) and <i><a href="/wiki/Epsilonretrovirus" title="Epsilonretrovirus">Epsilonretroviruses</a></i> (Type E). Families belonging to <b>Class II</b> show homology to mammalian <i><a href="/wiki/Betaretrovirus" title="Betaretrovirus">Betaretroviruses</a></i> (Type B) and <i><a href="/wiki/Deltaretrovirus" title="Deltaretrovirus">Deltaretroviruses</a></i> (Type D). Families belonging to <b>Class III</b> are similar to <a href="/wiki/Spumavirus" class="mw-redirect" title="Spumavirus">foamy viruses</a>. For all classes, if homologies appear well conserved in the <i>gag</i>, <i>pol</i>, and <i>env</i> gene, they are grouped into a <a href="/wiki/Superfamily_(molecular_biology)" class="mw-redirect" title="Superfamily (molecular biology)">superfamily</a>. There are more Class I families known to exist.<sup id="cite_ref-Belshaw2004_1-6" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid26800882_11-1" class="reference"><a href="#cite_note-pmid26800882-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The families themselves are named in a less uniform manner, with a mixture of naming based on an exogenous retrovirus, the priming tRNA (<a href="/wiki/Human_endogenous_retrovirus-W" title="Human endogenous retrovirus-W">HERV-W</a>, <a href="/wiki/Human_endogenous_retrovirus_K" title="Human endogenous retrovirus K">HERV-K</a>), or some neighboring gene (HERV-ADP), clone number (HERV-S71), or some amino acid motif (HERV-FRD). A proposed nomenclature aims to clean up the sometimes paraphyletic standards.<sup id="cite_ref-Nomenclature_for_endogenous_retrovi_6-1" class="reference"><a href="#cite_note-Nomenclature_for_endogenous_retrovi-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Origin">Origin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=11" title="Edit section: Origin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sometime during human evolution, exogenous progenitors of HERV inserted themselves into germ line cells and then replicated along with the host's genes using and exploiting the host's cellular mechanisms. Because of their distinct genomic structure, HERVs were subjected to many rounds of amplification and transposition, which led to a more widespread distribution of retroviral DNA.<sup id="cite_ref-Belshaw2004_1-7" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Nevertheless, one family of viruses has been active since the divergence of <a href="/wiki/Human" title="Human">humans</a> and <a href="/wiki/Common_chimpanzee" class="mw-redirect" title="Common chimpanzee">chimpanzees</a>. This family, termed <a href="/wiki/HERV-K" class="mw-redirect" title="HERV-K">HERV-K</a> (HML2), makes up less than 1% of HERV elements but is one of the most studied. There are indications it has even been active in the past few hundred thousand years, e.g., some human individuals carry more copies of HML2 than others.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Traditionally, age estimates of HERVs are performed by comparing the 5' and 3' <a href="/wiki/Long_terminal_repeat" title="Long terminal repeat">LTR</a> of a HERV; however, this method is only relevant for full-length HERVs. A recent method, called cross-sectional dating,<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> uses variations within a single LTR to estimate the ages of HERV insertions. This method is more precise in estimating HERV ages and can be used for any HERV insertions. Cross-sectional dating has been used to suggest that two members of HERV-K (HML2), HERV-K106 and HERV-K116, were active in the last 800,000 years and that HERV-K106 may have infected modern humans 150,000 years ago.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> However, the absence of known infectious members of the HERV-K (HML2) family, and the lack of elements with a full coding potential within the published human genome sequence, suggests to some that the family is less likely to be active at present. In 2006 and 2007, researchers working independently in France and USA recreated functional versions of HERV-K (HML2).<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Expression_of_HERV_proteins">Expression of HERV proteins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=12" title="Edit section: Expression of HERV proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The expression of HERV-K, a biologically active family of HERV, produces proteins found in placenta. Furthermore, the expression of the envelope genes of <a href="/wiki/HERV-W" class="mw-redirect" title="HERV-W">HERV-W</a> (<a rel="nofollow" class="external text" href="https://www.genenames.org/data/hgnc_data.php?hgnc_id=13525">ERVW-1</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20130919043913/http://www.genenames.org/data/hgnc_data.php?hgnc_id=13525">Archived</a> 2013-09-19 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a>) and <a href="/wiki/HERV-FRD" class="mw-redirect" title="HERV-FRD">HERV-FRD</a> (<a rel="nofollow" class="external text" href="https://www.genenames.org/data/hgnc_data.php?hgnc_id=33823">ERVFRD-1</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20121026003119/http://www.genenames.org/data/hgnc_data.php?hgnc_id=33823">Archived</a> 2012-10-26 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a>) produces syncytins which are important for the generation of the <a href="/wiki/Syncytiotrophoblast" title="Syncytiotrophoblast">syncytiotrophoblast</a> cell layer during placentogenesis by inducing cell-cell fusion.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/HUGO_Gene_Nomenclature_Committee" title="HUGO Gene Nomenclature Committee">HUGO Gene Nomenclature Committee</a> (HGNC) approves gene symbols for transcribed human ERVs.<sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Functional_impact">Functional impact</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=13" title="Edit section: Functional impact"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>MER41.AIM2 is an HERV that regulates the transcription of AIM2 (Absent in Melanoma 2) which encodes for a sensor of foreign cytosolic DNA. This acts as a binding site for AIM2, meaning that it is necessary for the transcription of AIM2. Researchers had shown this by deleting MER41.AIM2 in HeLa cells using CRISPR/Cas9, leading to an undetectable transcript level of AIM2 in modified HeLa cells. The control cells, which still contained the MER41.AIM2 ERV, were observed with normal amounts of AIM2 transcript. In terms of immunity, researchers concluded that MER41.AIM2 is necessary for an inflammatory response to infection.<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Activation_by_exogenous_viruses">Activation by exogenous viruses</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=14" title="Edit section: Activation by exogenous viruses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Considerable evidence indicates that HERVs can be reactivated by viral infections, such as: </p> <ul><li>retroviruses – <a href="/wiki/Human_immunodeficiency_virus_type_1" class="mw-redirect" title="Human immunodeficiency virus type 1">human immunodeficiency virus type-1</a> (<a href="/wiki/HIV-1" class="mw-redirect" title="HIV-1">HIV-1</a>), <a href="/wiki/Human_T-lymphotropic_virus_1" title="Human T-lymphotropic virus 1">human T-lymphotropic virus 1</a> (HTLV-1);</li> <li>RNA viruses – <a href="/wiki/Influenza_A_virus" title="Influenza A virus">influenza A virus</a>, <a href="/wiki/Hepatitis_C_virus" title="Hepatitis C virus">hepatitis C virus</a> (HCV), severe acute respiratory syndrome coronavirus-2 (<a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARSCoV-2</a>);</li> <li>DNA viruses – <a href="/wiki/Herpes_simplex_virus_type_1" class="mw-redirect" title="Herpes simplex virus type 1">herpes simplex virus type-1</a> (HSV-1), <a href="/wiki/Epstein%E2%80%93Barr_virus" title="Epstein–Barr virus">Epstein-Barr virus</a> (EBV), <a href="/wiki/Human_cytomegalovirus" class="mw-redirect" title="Human cytomegalovirus">human cytomegalovirus</a> (CMV), <a href="/wiki/Kaposi%27s_sarcoma-associated_herpesvirus" title="Kaposi's sarcoma-associated herpesvirus">Kaposi’s sarcoma-associated herpesvirus</a> (KSHV) <sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup></li></ul> <p>Several studies have shown that EBV is able to transactivate the expression of the normally inactive HERV-K18 Env protein, e.g., interacting with resting <a href="/wiki/B_cell" title="B cell">B cells</a> via the <a href="/wiki/CD21" class="mw-redirect" title="CD21">CD21</a> receptor. Further studies revealed that the mechanism of transactivation depends on the expression of the major EBV late gene transactivator, <a href="/wiki/EBNA-2" class="mw-redirect" title="EBNA-2">EBNA-2</a>. In-depth analysis completed the picture identifying the EBV latent membrane protein <a href="/wiki/Epstein%E2%80%93Barr_virus_latent_membrane_protein_2" title="Epstein–Barr virus latent membrane protein 2">LMP-2A</a> as a strong candidate for the HERV-K18 transactivation. HERV-K18 has also been reported to have superantigen activity (i.e., polyclonal T and B cell activation regardless of the specificity of their antigen receptor).<sup id="cite_ref-:1_69-0" class="reference"><a href="#cite_note-:1-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p><p>It has also been shown that in vitro binding of the EBV gp350 protein caused activation of MSRV<i>env</i> and syncytin-1 in B-cells, monocytes, macrophages, and astrocytes - cells that are involved in pathogenesis of <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a>.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> Monocytes, especially after their differentiation into macrophages, appeared to be the most responsive to EBVgp350, expressing even higher levels of HERV-W<i>env</i> than B cells. This finding is concordant with another study, which demonstrated that during infectious mononucleosis, EBV promoted the strongest activation of HERV-W/MSRV expression in monocytes compared to other blood cell types.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Immune_response_to_HERVs">Immune response to HERVs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=15" title="Edit section: Immune response to HERVs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Despite having been integrated into genomes of vertebrates for millions of years, ERVs represent an intermediate stage between exogenous viruses and the host genome; it is suggested that immunological tolerance to HERV-derived proteins and peptides is imperfect due to the epigenetic silencing of HERV in the thymus and bone marrow, which prevents deletion of all HERV-specific T and B cells.<sup id="cite_ref-:0_72-0" class="reference"><a href="#cite_note-:0-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> As evidence of this, immunization of non-human primates with ERV-derived antigens mounted robust polyfunctional <a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">cytotoxic T cell</a> response as well as high antibody titers. According to phylogenetic studies, among 30 HERV families existing in the human genome, <a href="/wiki/HERV-K" class="mw-redirect" title="HERV-K">HERV-K</a> (HML-2) elements which integrated most recently are the most intact and biologically active forms.<sup id="cite_ref-:1_69-1" class="reference"><a href="#cite_note-:1-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> HERV-K <i>env</i> and HERV-H <i>env</i>, considered to be a new class of tumor-associated antigens, have been found to promote strong cytotoxic T-cell responses in patients with various types of cancers.<sup id="cite_ref-:0_72-1" class="reference"><a href="#cite_note-:0-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> </p><p>On a level of the innate immune sensing of nucleic acids, single-stranded RNA (<a href="/wiki/SsRNA" class="mw-redirect" title="SsRNA">ssRNA</a>) and double-stranded RNA </p><p>(<a href="/wiki/Dsrna" class="mw-redirect" title="Dsrna">dsRNA</a>) derived from endogenous retroviruses are recognized by <a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern recognition receptors</a> (PRRs). </p><p><a href="/wiki/SsRNA" class="mw-redirect" title="SsRNA">SsRNAs</a> can be sensed by <a href="/wiki/Toll-like_receptor" title="Toll-like receptor">Toll-like receptors</a> <a href="/wiki/TLR_7" class="mw-redirect" title="TLR 7">TLR-7</a> and <a href="/wiki/TLR_8" class="mw-redirect" title="TLR 8">TLR-8</a>, resulting in secretion of <a href="/wiki/IFN-%CE%B1" class="mw-redirect" title="IFN-α">IFN-α</a> by stimulated <a href="/wiki/Dendritic_cell" title="Dendritic cell">dendritic cells</a> (DCs) and <a href="/wiki/Macrophage" title="Macrophage">macrophages</a>, which was observed for ssRNAs derived from <a href="/wiki/HIV-1" class="mw-redirect" title="HIV-1">HIV-1</a>.<sup id="cite_ref-:2_75-0" class="reference"><a href="#cite_note-:2-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Dsrna" class="mw-redirect" title="Dsrna">DsRNAs</a> might be one of the most immunogenic nucleic acid <a href="/wiki/Pathogen-associated_molecular_pattern" title="Pathogen-associated molecular pattern">pathogen-associated molecular patterns</a> (PAMPs), since they are not found in cells in a normal state. HERV-derived dsRNA can be recognized by <a href="/wiki/TLR_3" class="mw-redirect" title="TLR 3">TLR-3</a>, <a href="/wiki/RIG-I" title="RIG-I">RIG-I</a>, and <a href="/wiki/MDA5" title="MDA5">MDA5</a>; RIG-I and MDA5 are known to induce a <a href="/wiki/Interferon_type_I" title="Interferon type I">type I IFN</a> response.<sup id="cite_ref-:2_75-1" class="reference"><a href="#cite_note-:2-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p>When retrotranscribed into DNA, retroviruses can be sensed by cyclic guanosine monophosphate (GMP)-adenosine monophosphate (AMP) <a href="/wiki/CGAS%E2%80%93STING_cytosolic_DNA_sensing_pathway" title="CGAS–STING cytosolic DNA sensing pathway">synthase-stimulator of interferon genes (cGAS-STING) pathway</a>, leading to the activation of <a href="/wiki/NF-%CE%BAB" title="NF-κB">nuclear factor-kappa B</a> (NF-kB) and IFN regulatory factor 3 (IRF3), which in turn trigger a type I IFN response. DsDNA could also be sensed by DNA-dependent activator of IFN-regulatory factors (DAI); DNA:RNA hybrids could be recognized by <a href="/wiki/TLR_9" class="mw-redirect" title="TLR 9">TLR-9</a><sup id="cite_ref-:2_75-2" class="reference"><a href="#cite_note-:2-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p><p>The recognition of nucleic acids through PRRs provides a very efficient strategy to fight against viral infections, at the same time exposing the host to a risk due to the possibility of recognizing self-nucleic acids and promotion of autoimmunity.<sup id="cite_ref-:2_75-3" class="reference"><a href="#cite_note-:2-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> Not surprisingly, HERVs have been found to be associated with different autoimmune and inflammatory diseases, such as <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a>, <a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">amyotrophic lateral sclerosis (ALS)</a>, <a href="/wiki/Lupus" title="Lupus">systemic lupus erythematosus (SLE)</a>, <a href="/wiki/Rheumatoid_arthritis" title="Rheumatoid arthritis">rheumathoid arthritis (RA)</a>, and <a href="/wiki/Sj%C3%B6gren_syndrome" title="Sjögren syndrome">Sjögren syndrome (SS)</a>.<sup id="cite_ref-:1_69-2" class="reference"><a href="#cite_note-:1-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p><p>On a protein level, a direct interaction between TLRs and certain HERV proteins has been shown. For example, the surface unit of HERV-W Env (also known as multiple sclerosis-associated retroviral element (MSRV) env) was found to bind to <a href="/wiki/Toll-like_receptor_4" title="Toll-like receptor 4">TLR4</a> and <a href="/wiki/CD14" title="CD14">CD14</a>, stimulating the production of pro-inflammatory cytokines including <a href="/wiki/Interleukin_1_beta" title="Interleukin 1 beta">IL-1β</a>, <a href="/wiki/Interleukin_6" title="Interleukin 6">IL-6</a>, and <a href="/wiki/TNF%CE%B1" class="mw-redirect" title="TNFα">TNFα</a>. HERV-W Env can trigger a maturation process in human <a href="/wiki/Dendritic_cell" title="Dendritic cell">dendritic cells</a>, endowing them with the capacity to support a <a href="/wiki/Th1_cell" class="mw-redirect" title="Th1 cell">Th1</a>-like type of Th cell differentiation.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> </p><p>Immunological studies have shown some evidence for <a href="/wiki/T_cell" title="T cell">T cell</a> immune responses against HERVs in HIV-infected individuals.<sup id="cite_ref-garrison_78-0" class="reference"><a href="#cite_note-garrison-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> The hypothesis that HIV induces HERV expression in HIV-infected cells led to the proposal that a vaccine targeting HERV antigens could specifically eliminate HIV-infected cells. The potential advantage of this novel approach is that, by using HERV antigens as surrogate markers of HIV-infected cells, it could circumvent the difficulty inherent in directly targeting notoriously diverse and fast-mutating HIV antigens.<sup id="cite_ref-garrison_78-1" class="reference"><a href="#cite_note-garrison-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Techniques_for_characterizing_ERVs">Techniques for characterizing ERVs</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=16" title="Edit section: Techniques for characterizing ERVs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Whole_genome_sequencing">Whole genome sequencing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=17" title="Edit section: Whole genome sequencing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Example: A PERV Chinese-born <a href="/wiki/Minipig" class="mw-redirect" title="Minipig">minipig</a> isolate, PERV-A-BM, was sequenced completely and along with different breeds and cell lines in order to understand its genetic variation and evolution. The observed number of nucleotide substitutions and among the different genome sequences helped researchers determine an estimated age that PERV-A-BM was integrated into its host genome, which was found to be of an evolutionary age earlier than the European-born pigs isolates.<sup id="cite_ref-Tang16_54-1" class="reference"><a href="#cite_note-Tang16-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Chromatin_immunoprecipitation_with_sequencing_(ChIP-seq)"><span id="Chromatin_immunoprecipitation_with_sequencing_.28ChIP-seq.29"></span>Chromatin immunoprecipitation with sequencing (ChIP-seq)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=18" title="Edit section: Chromatin immunoprecipitation with sequencing (ChIP-seq)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This technique is used to find histone marks indicative of promoters and enhancers, which are binding sites for DNA proteins, and repressed regions and trimethylation.<sup id="cite_ref-Chuong_325–329_34-1" class="reference"><a href="#cite_note-Chuong_325–329-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> DNA methylation has been shown to be vital to maintain silencing of ERVs in mouse somatic cells, while histone marks are vital for the same purpose in <a href="/wiki/Embryonic_stem_cells" class="mw-redirect" title="Embryonic stem cells">embryonic stem cells</a> (ESCs) and early embryogenesis.<sup id="cite_ref-Rebollo12_7-2" class="reference"><a href="#cite_note-Rebollo12-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=19" title="Edit section: Applications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Constructing_phylogenies">Constructing phylogenies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=20" title="Edit section: Constructing phylogenies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Because most HERVs have no function, are selectively neutral, and are very abundant in primate genomes, they easily serve as phylogenetic markers for linkage analysis. They can be exploited by comparing the integration site polymorphisms or the evolving, proviral, nucleotide sequences of orthologs. To estimate when integration occurred, researchers used distances from each phylogenetic tree to find the rate of molecular evolution at each particular locus. It is also useful that ERVs are abundant within the genomes of many species (i.e., plants, insects, mollusks, fish, rodents, domestic pets, and livestock) and the application of this ERV approach can be used to answer a variety of phylogenetic questions.<sup id="cite_ref-Johnson_10254–10260_9-2" class="reference"><a href="#cite_note-Johnson_10254–10260-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Designating_the_age_of_provirus_and_the_time_points_of_species_separation_events">Designating the age of provirus and the time points of species separation events</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=21" title="Edit section: Designating the age of provirus and the time points of species separation events"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This is accomplished by comparing the different HERV from different evolutionary periods. For example, this study was done for different hominoids, which ranged from humans to apes and to monkeys. This is difficult to do with PERV because of the large diversity present.<sup id="cite_ref-Tönjes03_55-1" class="reference"><a href="#cite_note-Tönjes03-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Further_research">Further research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=22" title="Edit section: Further research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Epigenetic_variability">Epigenetic variability</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=23" title="Edit section: Epigenetic variability"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Researchers could analyze individual epigenomes and <a href="/wiki/Transcriptomes" class="mw-redirect" title="Transcriptomes">transcriptomes</a> to study the reactivation of dormant transposable elements through epigenetic release and their potential associations with human disease and exploring the specifics of gene regulatory networks.<sup id="cite_ref-Rebollo12_7-3" class="reference"><a href="#cite_note-Rebollo12-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Immunological_problems_of_xenotransplantation">Immunological problems of xenotransplantation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=24" title="Edit section: Immunological problems of xenotransplantation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Little is known about an effective way to overcoming <a href="/wiki/Hyperacute_rejection" class="mw-redirect" title="Hyperacute rejection">hyperacute rejection</a> (HAR), which follows the activation of complement initiated by xenoreactive antibodies recognizing galactosyl-alpha1-3galatosyl (alpha-Gal) antigens on the donor epithelium.<sup id="cite_ref-Patience97_50-3" class="reference"><a href="#cite_note-Patience97-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Risk_factors_of_HERVs_in_gene_therapy">Risk factors of HERVs in gene therapy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=25" title="Edit section: Risk factors of HERVs in gene therapy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Because retroviruses are able to recombine with each other and with other endogenous DNA sequences, it would be beneficial for gene therapy to explore the potential risks HERVs can cause, if any. Also, this ability of HERVs to recombine can be manipulated for site-directed integration by including HERV sequences in retroviral vectors.<sup id="cite_ref-Belshaw2004_1-8" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="HERV_gene_expression">HERV gene expression</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=26" title="Edit section: HERV gene expression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Researchers believe that RNA and proteins encoded for by HERV genes should continue to be explored for putative function in cell physiology and in pathological conditions. These studies would examine and more deeply define the biological significance of the proteins synthesized.<sup id="cite_ref-Belshaw2004_1-9" class="reference"><a href="#cite_note-Belshaw2004-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=27" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1259569809">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{clear:left;float:left;margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><span class="portalbox-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Sida-aids.png" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/28px-Sida-aids.png" decoding="async" width="28" height="28" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/42px-Sida-aids.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/56px-Sida-aids.png 2x" data-file-width="290" data-file-height="290" /></a></span></span><span class="portalbox-link"><a href="/wiki/Portal:Viruses" title="Portal:Viruses">Viruses portal</a></span></li></ul> <ul><li><a href="/wiki/Avian_sarcoma_leukosis_virus" title="Avian sarcoma leukosis virus">Avian sarcoma leukosis virus</a> (ASLV)</li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Endogeny_(biology)" title="Endogeny (biology)">Endogeny (biology)</a></li> <li><a href="/wiki/ERV3" title="ERV3">ERV3</a></li> <li><a href="/wiki/HERV-FRD" class="mw-redirect" title="HERV-FRD">HERV-FRD</a></li> <li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Jaagsiekte_sheep_retrovirus" title="Jaagsiekte sheep retrovirus">Jaagsiekte sheep retrovirus</a> (JSRV)</li> <li><a href="/wiki/Koala_retrovirus" title="Koala retrovirus">Koala retrovirus</a> (KoRV)</li> <li><a href="/wiki/Mouse_mammary_tumor_virus" title="Mouse mammary tumor virus">Mouse mammary tumor virus</a> (MMTV)</li> <li><a href="/wiki/Murine_leukemia_virus" title="Murine leukemia virus">Murine leukemia virus</a> (MLV) and <a href="/wiki/Xenotropic_murine_leukemia_virus-related_virus" class="mw-redirect" title="Xenotropic murine leukemia virus-related virus">xenotropic murine leukemia virus-related virus</a> (XMRV)</li> <li><a href="/wiki/Paleovirology" title="Paleovirology">Paleovirology</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=28" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Belshaw2004-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Belshaw2004_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Belshaw2004_1-9"><sup><i><b>j</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBelshawPereiraKatzourakisTalbot2004" class="citation journal cs1">Belshaw R, Pereira V, Katzourakis A, Talbot G, Paces J, Burt A, Tristem M (April 2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC387345">"Long-term reinfection of the human genome by endogenous retroviruses"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>101</b> (14): 4894–4899. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2004PNAS..101.4894B">2004PNAS..101.4894B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Virology&rft.atitle=Emergence+of+vertebrate+retroviruses+and+envelope+capture&rft.volume=318&rft.issue=1&rft.pages=183-191&rft.date=2004-01&rft_id=info%3Adoi%2F10.1016%2Fj.virol.2003.09.026&rft_id=info%3Apmid%2F14972546&rft.aulast=Kim&rft.aufirst=FJ&rft.au=Battini%2C+JL&rft.au=Manel%2C+N&rft.au=Sitbon%2C+M&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.virol.2003.09.026&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStoyeBoeke1997" class="citation encyclopaedia cs1">Stoye JP, Boeke JD (1997). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK19468/">"Retrotransposons, Endogenous 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Cold Spring Harbor (NY): Cold Spring Harbor Laboratory Press. p. 343. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780879695712" title="Special:BookSources/9780879695712"><bdi>9780879695712</bdi></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/21433351">21433351</a>. NBK19468<span class="reference-accessdate">. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cancers&rft.atitle=Human+Endogenous+Retroviruses+%28HERVs%29%3A+Shaping+the+Innate+Immune+Response+in+Cancers&rft.volume=12&rft.issue=3&rft.pages=610&rft.date=2020-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7139688%23id-name%3DPMC&rft_id=info%3Apmid%2F32155827&rft_id=info%3Adoi%2F10.3390%2Fcancers12030610&rft.aulast=Alcazer&rft.aufirst=V&rft.au=Bonaventura%2C+P&rft.au=Depil%2C+S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7139688&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></span> </li> <li id="cite_note-76"><span class="mw-cite-backlink"><b><a href="#cite_ref-76">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHurstMagiorkinis2015" class="citation journal cs1">Hurst TP, Magiorkinis G (June 2015). <a rel="nofollow" class="external text" href="https://doi.org/10.1099%2Fjgv.0.000017">"Activation of the innate immune response by endogenous retroviruses"</a>. <i>The Journal of General Virology</i>. <b>96</b> (Pt 6): 1207–1218. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1099%2Fjgv.0.000017">10.1099/jgv.0.000017</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26068187">26068187</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+General+Virology&rft.atitle=Activation+of+the+innate+immune+response+by+endogenous+retroviruses&rft.volume=96&rft.issue=Pt+6&rft.pages=1207-1218&rft.date=2015-06&rft_id=info%3Adoi%2F10.1099%2Fjgv.0.000017&rft_id=info%3Apmid%2F26068187&rft.aulast=Hurst&rft.aufirst=TP&rft.au=Magiorkinis%2C+G&rft_id=https%3A%2F%2Fdoi.org%2F10.1099%252Fjgv.0.000017&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></span> </li> <li id="cite_note-77"><span class="mw-cite-backlink"><b><a href="#cite_ref-77">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRollandJouvin-MarcheViretFaure2006" class="citation journal cs1">Rolland A, Jouvin-Marche E, Viret C, Faure M, Perron H, Marche PN (June 2006). <a rel="nofollow" class="external text" href="https://doi.org/10.4049%2Fjimmunol.176.12.7636">"The envelope protein of a human endogenous retrovirus-W family activates innate immunity through CD14/TLR4 and promotes Th1-like responses"</a>. <i>Journal of Immunology</i>. <b>176</b> (12): 7636–7644. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.4049%2Fjimmunol.176.12.7636">10.4049/jimmunol.176.12.7636</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16751411">16751411</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:24492010">24492010</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Immunology&rft.atitle=The+envelope+protein+of+a+human+endogenous+retrovirus-W+family+activates+innate+immunity+through+CD14%2FTLR4+and+promotes+Th1-like+responses&rft.volume=176&rft.issue=12&rft.pages=7636-7644&rft.date=2006-06&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A24492010%23id-name%3DS2CID&rft_id=info%3Apmid%2F16751411&rft_id=info%3Adoi%2F10.4049%2Fjimmunol.176.12.7636&rft.aulast=Rolland&rft.aufirst=A&rft.au=Jouvin-Marche%2C+E&rft.au=Viret%2C+C&rft.au=Faure%2C+M&rft.au=Perron%2C+H&rft.au=Marche%2C+PN&rft_id=https%3A%2F%2Fdoi.org%2F10.4049%252Fjimmunol.176.12.7636&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></span> </li> <li id="cite_note-garrison-78"><span class="mw-cite-backlink">^ <a href="#cite_ref-garrison_78-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-garrison_78-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGarrisonJonesMeiklejohnAnwar2007" class="citation journal cs1">Garrison KE, Jones RB, Meiklejohn DA, Anwar N, Ndhlovu LC, Chapman JM, et al. (November 2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2065876">"T cell responses to human endogenous retroviruses in HIV-1 infection"</a>. <i>PLOS Pathogens</i>. <b>3</b> (11): e165. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.ppat.0030165">10.1371/journal.ppat.0030165</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2065876">2065876</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17997601">17997601</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+Pathogens&rft.atitle=T+cell+responses+to+human+endogenous+retroviruses+in+HIV-1+infection&rft.volume=3&rft.issue=11&rft.pages=e165&rft.date=2007-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2065876%23id-name%3DPMC&rft_id=info%3Apmid%2F17997601&rft_id=info%3Adoi%2F10.1371%2Fjournal.ppat.0030165&rft.aulast=Garrison&rft.aufirst=KE&rft.au=Jones%2C+RB&rft.au=Meiklejohn%2C+DA&rft.au=Anwar%2C+N&rft.au=Ndhlovu%2C+LC&rft.au=Chapman%2C+JM&rft.au=Erickson%2C+AL&rft.au=Agrawal%2C+A&rft.au=Spotts%2C+G&rft.au=Hecht%2C+FM&rft.au=Rakoff-Nahoum%2C+S&rft.au=Lenz%2C+J&rft.au=Ostrowski%2C+MA&rft.au=Nixon%2C+DF&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2065876&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=29" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLöwerLöwerKurth1996" class="citation journal cs1">Löwer R, Löwer J, Kurth R (May 1996). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC39218">"The viruses in all of us: characteristics and biological significance of human endogenous retrovirus sequences"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>93</b> (11): 5177–5184. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1996PNAS...93.5177L">1996PNAS...93.5177L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.93.11.5177">10.1073/pnas.93.11.5177</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC39218">39218</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/8643549">8643549</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=The+viruses+in+all+of+us%3A+characteristics+and+biological+significance+of+human+endogenous+retrovirus+sequences&rft.volume=93&rft.issue=11&rft.pages=5177-5184&rft.date=1996-05&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC39218%23id-name%3DPMC&rft_id=info%3Apmid%2F8643549&rft_id=info%3Adoi%2F10.1073%2Fpnas.93.11.5177&rft_id=info%3Abibcode%2F1996PNAS...93.5177L&rft.aulast=L%C3%B6wer&rft.aufirst=R&rft.au=L%C3%B6wer%2C+J&rft.au=Kurth%2C+R&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC39218&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMolèsTesniereGuilhou2005" class="citation journal cs1">Molès JP, Tesniere A, Guilhou JJ (July 2005). "A new endogenous retroviral sequence is expressed in skin of patients with psoriasis". <i>The British Journal of Dermatology</i>. <b>153</b> (1): 83–89. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2133.2005.06555.x">10.1111/j.1365-2133.2005.06555.x</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/16029331">16029331</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:6642536">6642536</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+British+Journal+of+Dermatology&rft.atitle=A+new+endogenous+retroviral+sequence+is+expressed+in+skin+of+patients+with+psoriasis&rft.volume=153&rft.issue=1&rft.pages=83-89&rft.date=2005-07&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A6642536%23id-name%3DS2CID&rft_id=info%3Apmid%2F16029331&rft_id=info%3Adoi%2F10.1111%2Fj.1365-2133.2005.06555.x&rft.aulast=Mol%C3%A8s&rft.aufirst=JP&rft.au=Tesniere%2C+A&rft.au=Guilhou%2C+JJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSeifarthFrankZeilfelderSpiess2005" class="citation journal cs1">Seifarth W, Frank O, Zeilfelder U, Spiess B, Greenwood AD, Hehlmann R, Leib-Mösch C (January 2005). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC538696">"Comprehensive analysis of human endogenous retrovirus transcriptional activity in human tissues with a retrovirus-specific microarray"</a>. <i>Journal of Virology</i>. <b>79</b> (1): 341–352. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1128%2FJVI.79.1.341-352.2005">10.1128/JVI.79.1.341-352.2005</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC538696">538696</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15596828">15596828</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Virology&rft.atitle=Comprehensive+analysis+of+human+endogenous+retrovirus+transcriptional+activity+in+human+tissues+with+a+retrovirus-specific+microarray&rft.volume=79&rft.issue=1&rft.pages=341-352&rft.date=2005-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC538696%23id-name%3DPMC&rft_id=info%3Apmid%2F15596828&rft_id=info%3Adoi%2F10.1128%2FJVI.79.1.341-352.2005&rft.aulast=Seifarth&rft.aufirst=W&rft.au=Frank%2C+O&rft.au=Zeilfelder%2C+U&rft.au=Spiess%2C+B&rft.au=Greenwood%2C+AD&rft.au=Hehlmann%2C+R&rft.au=Leib-M%C3%B6sch%2C+C&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC538696&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKnerrBeinderRascher2002" class="citation journal cs1">Knerr I, Beinder E, Rascher W (February 2002). "Syncytin, a novel human endogenous retroviral gene in human placenta: evidence for its dysregulation in preeclampsia and HELLP syndrome". <i>American Journal of Obstetrics and Gynecology</i>. <b>186</b> (2): 210–213. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1067%2Fmob.2002.119636">10.1067/mob.2002.119636</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/11854637">11854637</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+Obstetrics+and+Gynecology&rft.atitle=Syncytin%2C+a+novel+human+endogenous+retroviral+gene+in+human+placenta%3A+evidence+for+its+dysregulation+in+preeclampsia+and+HELLP+syndrome&rft.volume=186&rft.issue=2&rft.pages=210-213&rft.date=2002-02&rft_id=info%3Adoi%2F10.1067%2Fmob.2002.119636&rft_id=info%3Apmid%2F11854637&rft.aulast=Knerr&rft.aufirst=I&rft.au=Beinder%2C+E&rft.au=Rascher%2C+W&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGiffordTristem2003" class="citation journal cs1">Gifford R, Tristem M (May 2003). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20130215200652/http://www.vgb.ucl.ac.uk/pdf/Gifford_2003.pdf">"The evolution, distribution and diversity of endogenous retroviruses"</a> <span class="cs1-format">(PDF)</span>. <i>Virus Genes</i>. <b>26</b> (3): 291–315. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1023%2FA%3A1024455415443">10.1023/A:1024455415443</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/12876457">12876457</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:34639116">34639116</a>. Archived from <a rel="nofollow" class="external text" href="http://www.vgb.ucl.ac.uk/pdf/Gifford_2003.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 2013-02-15<span class="reference-accessdate">. Retrieved <span class="nowrap">2008-01-16</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Virus+Genes&rft.atitle=The+evolution%2C+distribution+and+diversity+of+endogenous+retroviruses&rft.volume=26&rft.issue=3&rft.pages=291-315&rft.date=2003-05&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A34639116%23id-name%3DS2CID&rft_id=info%3Apmid%2F12876457&rft_id=info%3Adoi%2F10.1023%2FA%3A1024455415443&rft.aulast=Gifford&rft.aufirst=R&rft.au=Tristem%2C+M&rft_id=http%3A%2F%2Fwww.vgb.ucl.ac.uk%2Fpdf%2FGifford_2003.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndogenous+retrovirus" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Endogenous_retrovirus&action=edit&section=30" title="Edit section: External 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<ul><li><a href="/wiki/Ground_squirrel_hepatitis_virus" title="Ground squirrel hepatitis virus">Ground squirrel hepatitis virus</a></li> <li><a href="/wiki/Hepatitis_B_virus" title="Hepatitis B virus">Hepatitis B virus</a></li> <li><a href="/wiki/Woodchuck_hepatitis_virus" title="Woodchuck hepatitis virus">Woodchuck hepatitis virus</a></li> <li><a href="/wiki/Woolly_monkey_hepatitis_B_virus" title="Woolly monkey hepatitis B virus">Woolly monkey hepatitis B virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Parahepadnavirus&action=edit&redlink=1" class="new" title="Parahepadnavirus (page does not exist)">Parahepadnavirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Ortervirales" title="Ortervirales">Ortervirales</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">(<a href="/wiki/SsRNA-RT_virus" class="mw-redirect" title="SsRNA-RT virus">ssRNA-RT viruses</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Belpaoviridae" class="mw-redirect" title="Belpaoviridae">Belpaoviridae</a></i></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 id="Semotivirus" scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Semotivirus" title="Semotivirus">Semotivirus</a></i></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/Caenorhabditis_elegans_Cer13_virus" title="Caenorhabditis elegans Cer13 virus">Caenorhabditis elegans Cer13 virus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Metaviridae" title="Metaviridae">Metaviridae</a></i></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%"><i><a href="/w/index.php?title=Errantivirus&action=edit&redlink=1" class="new" title="Errantivirus (page does not exist)">Errantivirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Metavirus" title="Metavirus">Metavirus</a></i></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/Caenorhabditis_elegans_Cer1_virus" title="Caenorhabditis elegans Cer1 virus">Caenorhabditis elegans Cer1 virus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Pseudoviridae" title="Pseudoviridae">Pseudoviridae</a></i></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%"><i><a href="/w/index.php?title=Hemivirus&action=edit&redlink=1" class="new" title="Hemivirus (page does not exist)">Hemivirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Pseudovirus_(genus)" class="mw-redirect" title="Pseudovirus (genus)">Pseudovirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Sirevirus&action=edit&redlink=1" class="new" title="Sirevirus (page does not exist)">Sirevirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Retrovirus" title="Retrovirus">Retroviridae</a></i></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%"><i><a href="/wiki/Orthoretrovirinae" title="Orthoretrovirinae">Orthoretrovirinae</a></i></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%"><i><a href="/wiki/Alpharetrovirus" title="Alpharetrovirus">Alpharetrovirus</a></i></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/Avian_sarcoma_leukosis_virus" title="Avian sarcoma leukosis virus">Avian sarcoma leukosis virus</a></li> <li><a href="/wiki/Rous_sarcoma_virus" title="Rous sarcoma virus">Rous sarcoma virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Betaretrovirus" title="Betaretrovirus">Betaretrovirus</a></i></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/Mouse_mammary_tumor_virus" title="Mouse mammary tumor virus">Mouse mammary tumor virus</a></li> <li><a href="/wiki/Jaagsiekte_sheep_retrovirus" title="Jaagsiekte sheep retrovirus">Jaagsiekte sheep retrovirus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Deltaretrovirus" title="Deltaretrovirus">Deltaretrovirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_T-lymphotropic_virus" class="mw-redirect" title="Human T-lymphotropic virus">Human T-lymphotropic virus</a> (<a href="/wiki/Human_T-lymphotropic_virus_1" title="Human T-lymphotropic virus 1">HTLV-1</a></li> <li><a href="/wiki/Human_T-lymphotropic_virus_2" title="Human T-lymphotropic virus 2">HTLV-2</a>, 3, 4)</li> <li><a href="/wiki/Simian-T-lymphotropic_virus" class="mw-redirect" title="Simian-T-lymphotropic virus">Simian-T-lymphotropic virus</a> (types 1-4)</li> <li><a href="/wiki/Bovine_leukemia_virus" title="Bovine leukemia virus">Bovine leukemia virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Epsilonretrovirus" title="Epsilonretrovirus">Epsilonretrovirus</a></i></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/Walleye_epidermal_hyperplasia_virus" title="Walleye epidermal hyperplasia virus">Walleye epidermal hyperplasia virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Gammaretrovirus" title="Gammaretrovirus">Gammaretrovirus</a></i></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/Murine_leukemia_virus" title="Murine leukemia virus">Murine leukemia virus</a></li> <li><a href="/wiki/Abelson_murine_leukemia_virus" title="Abelson murine leukemia virus">Abelson murine leukemia virus</a></li> <li><a href="/wiki/Friend_virus" title="Friend virus">Friend virus</a></li> <li><a href="/wiki/Feline_leukemia_virus" title="Feline leukemia virus">Feline leukemia virus</a></li> <li><a href="/wiki/Koala_retrovirus" title="Koala retrovirus">Koala retrovirus</a> (KoRV)</li> <li><a href="/wiki/Xenotropic_murine_leukemia_virus-related_virus" class="mw-redirect" title="Xenotropic murine leukemia virus-related virus">Xenotropic murine leukemia virus-related virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Lentivirus" title="Lentivirus">Lentivirus</a></i></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/HIV" title="HIV">HIV</a> (human immunodeficiency viruses)</li> <li><a href="/wiki/Simian_immunodeficiency_virus" title="Simian immunodeficiency virus">Simian immunodeficiency viruses</a> (SIV)</li> <li><a href="/wiki/Feline_immunodeficiency_virus" title="Feline immunodeficiency virus">Feline immunodeficiency virus</a> (FIV)</li> <li><a href="/wiki/Puma_lentivirus" title="Puma lentivirus">Puma lentivirus</a> (PLV)</li> <li><a href="/wiki/Equine_infectious_anemia" title="Equine infectious anemia">Equine infectious anemia virus</a> (EIAV)</li> <li><a href="/wiki/Bovine_immunodeficiency_virus" title="Bovine immunodeficiency virus">Bovine immunodeficiency virus</a> (BIV)</li> <li><a href="/wiki/Caprine_arthritis_encephalitis_virus" title="Caprine arthritis encephalitis virus">Caprine arthritis encephalitis virus</a></li> <li><a href="/wiki/Visna-maedi_virus" title="Visna-maedi virus">Visna-maedi virus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Spumaretrovirinae" title="Spumaretrovirinae">Spumaretrovirinae</a></i></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%"><i><a href="/w/index.php?title=Bovispumavirus&action=edit&redlink=1" class="new" title="Bovispumavirus (page does not exist)">Bovispumavirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Equispumavirus&action=edit&redlink=1" class="new" title="Equispumavirus (page does not exist)">Equispumavirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Felispumavirus&action=edit&redlink=1" class="new" title="Felispumavirus (page does not exist)">Felispumavirus</a></i></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/Feline_foamy_virus" title="Feline foamy virus">Feline foamy virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Prosimiispumavirus&action=edit&redlink=1" class="new" title="Prosimiispumavirus (page does not exist)">Prosimiispumavirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Simiispumavirus&action=edit&redlink=1" class="new" title="Simiispumavirus (page does not exist)">Simiispumavirus</a></i></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/Simian_foamy_virus" class="mw-redirect" title="Simian foamy virus">Simian foamy virus</a></li> <li><a href="/wiki/Human_foamy_virus" title="Human foamy virus">Human foamy virus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">(<a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT viruses</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 id="Caulimoviridae" scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Caulimoviridae" title="Caulimoviridae">Caulimoviridae</a></i></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%"><i><a href="/wiki/Badnavirus" title="Badnavirus">Badnavirus</a></i></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/Banana_streak_MY_virus" title="Banana streak MY virus">Banana streak MY virus</a></li> <li><a href="/wiki/Cacao_swollen_shoot_virus" title="Cacao swollen shoot virus">Cacao swollen shoot virus</a></li> <li><a href="/wiki/Kalanchoe_top-spotting_virus" title="Kalanchoe top-spotting virus">Kalanchoe top-spotting virus</a></li> <li><a href="/wiki/Rubus_yellow_net_virus" title="Rubus yellow net virus">Rubus yellow net virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Caulimovirus" title="Caulimovirus">Caulimovirus</a></i></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/Carnation_etched_ring_virus" title="Carnation etched ring virus">Carnation etched ring virus</a></li> <li><a href="/wiki/Cauliflower_mosaic_virus" title="Cauliflower mosaic virus">Cauliflower mosaic virus</a></li> <li><a href="/wiki/Strawberry_vein_banding_virus" title="Strawberry vein banding virus">Strawberry vein banding virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Cavemovirus" title="Cavemovirus">Cavemovirus</a></i></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/Cassava_vein_mosaic_virus" title="Cassava vein mosaic virus">Cassava vein mosaic virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Dioscovirus&action=edit&redlink=1" class="new" title="Dioscovirus (page does not exist)">Dioscovirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Petuvirus" title="Petuvirus">Petuvirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Rosadnavirus&action=edit&redlink=1" class="new" title="Rosadnavirus (page does not exist)">Rosadnavirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Ruflodivirus&action=edit&redlink=1" class="new" title="Ruflodivirus (page does not exist)">Ruflodivirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Solendovirus" title="Solendovirus">Solendovirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Soymovirus" title="Soymovirus">Soymovirus</a></i></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/Soybean_chlorotic_mottle_virus" title="Soybean chlorotic mottle virus">Soybean chlorotic mottle virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Tungrovirus" title="Tungrovirus">Tungrovirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/w/index.php?title=Vaccinivirus&action=edit&redlink=1" class="new" title="Vaccinivirus (page does not exist)">Vaccinivirus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li class="mw-empty-elt"></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Endogenous</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/ERV-Fc" title="ERV-Fc">ERV-Fc</a></li> <li><a href="/wiki/Syncytin-1" title="Syncytin-1">ERVWE1</a></li> <li><a href="/wiki/HCP5" title="HCP5">HCP5</a></li> <li><a href="/wiki/Human_teratocarcinoma-derived_virus" class="mw-redirect" title="Human teratocarcinoma-derived virus">Human teratocarcinoma-derived virus</a></li> <li><a href="/wiki/Human_Endogenous_Retrovirus-W" class="mw-redirect" title="Human Endogenous Retrovirus-W">Human Endogenous Retrovirus-W</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Self-replicating_organic_structures" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align: center;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Self-replicating_organic_structures" title="Template:Self-replicating organic structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Self-replicating_organic_structures" title="Template talk:Self-replicating organic structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Self-replicating_organic_structures" title="Special:EditPage/Template:Self-replicating organic structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Self-replicating_organic_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br />agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus<br />dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br />self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br />elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a class="mw-selflink selflink">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Endogenous_retrovirus&oldid=1259796292">https://en.wikipedia.org/w/index.php?title=Endogenous_retrovirus&oldid=1259796292</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Molecular_biology" title="Category:Molecular biology">Molecular biology</a></li><li><a href="/wiki/Category:Endogenous_retroviruses" title="Category:Endogenous retroviruses">Endogenous retroviruses</a></li><li><a href="/wiki/Category:Swine_diseases" 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