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</ul> </li> <li id="toc-Modern_synthesis_and_its_successors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Modern_synthesis_and_its_successors"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Modern synthesis and its successors</span> </div> </a> <ul id="toc-Modern_synthesis_and_its_successors-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Molecular_basis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Molecular_basis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Molecular basis</span> </div> </a> <button aria-controls="toc-Molecular_basis-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 Molecular basis subsection</span> </button> <ul id="toc-Molecular_basis-sublist" class="vector-toc-list"> <li id="toc-DNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>DNA</span> </div> </a> <ul id="toc-DNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Chromosomes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Chromosomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Chromosomes</span> </div> </a> <ul id="toc-Chromosomes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Structure_and_function" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Structure_and_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Structure and function</span> </div> </a> <button aria-controls="toc-Structure_and_function-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 Structure and function subsection</span> </button> <ul id="toc-Structure_and_function-sublist" class="vector-toc-list"> <li id="toc-Structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Structure</span> </div> </a> <ul id="toc-Structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Complexity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Complexity"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Complexity</span> </div> </a> <ul id="toc-Complexity-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Gene_expression" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Gene_expression"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Gene expression</span> </div> </a> <button aria-controls="toc-Gene_expression-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 Gene expression subsection</span> </button> <ul id="toc-Gene_expression-sublist" class="vector-toc-list"> <li id="toc-Genetic_code" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_code"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Genetic code</span> </div> </a> <ul id="toc-Genetic_code-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transcription" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transcription"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Transcription</span> </div> </a> <ul id="toc-Transcription-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Translation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Translation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Translation</span> </div> </a> <ul id="toc-Translation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Regulation</span> </div> </a> <ul id="toc-Regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_genes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.5</span> <span>RNA genes</span> </div> </a> <ul id="toc-RNA_genes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Inheritance" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Inheritance"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Inheritance</span> </div> </a> <button aria-controls="toc-Inheritance-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 Inheritance subsection</span> </button> <ul id="toc-Inheritance-sublist" class="vector-toc-list"> <li id="toc-Mendelian_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mendelian_inheritance"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Mendelian inheritance</span> </div> </a> <ul id="toc-Mendelian_inheritance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_replication_and_cell_division" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_replication_and_cell_division"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>DNA replication and cell division</span> </div> </a> <ul id="toc-DNA_replication_and_cell_division-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Molecular_inheritance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Molecular_inheritance"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Molecular inheritance</span> </div> </a> <ul id="toc-Molecular_inheritance-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genome" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genome"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Genome</span> </div> </a> <button aria-controls="toc-Genome-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 Genome subsection</span> </button> <ul id="toc-Genome-sublist" class="vector-toc-list"> <li id="toc-Number_of_genes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Number_of_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Number of genes</span> </div> </a> <ul id="toc-Number_of_genes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Essential_genes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Essential_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Essential genes</span> </div> </a> <ul id="toc-Essential_genes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_and_genomic_nomenclature" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_and_genomic_nomenclature"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Genetic and genomic nomenclature</span> </div> </a> <ul id="toc-Genetic_and_genomic_nomenclature-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genetic_engineering" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genetic_engineering"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Genetic engineering</span> </div> </a> <ul id="toc-Genetic_engineering-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">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> <button aria-controls="toc-References-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 References subsection</span> </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Citations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Citations"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Citations</span> </div> </a> <ul id="toc-Citations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sources" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sources"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.2</span> <span>Sources</span> </div> </a> <ul id="toc-Sources-sublist" class="vector-toc-list"> </ul> </li> </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" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Gene</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 117 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-117" 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">117 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Geen_(genetika)" title="Geen (genetika) – Afrikaans" lang="af" hreflang="af" data-title="Geen (genetika)" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-als mw-list-item"><a href="https://als.wikipedia.org/wiki/Gen" title="Gen – Alemannic" lang="gsw" hreflang="gsw" data-title="Gen" data-language-autonym="Alemannisch" data-language-local-name="Alemannic" class="interlanguage-link-target"><span>Alemannisch</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AC%D9%8A%D9%86" title="جين – Arabic" lang="ar" hreflang="ar" data-title="جين" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Xen" title="Xen – Asturian" lang="ast" hreflang="ast" data-title="Xen" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Gen" title="Gen – Azerbaijani" lang="az" hreflang="az" data-title="Gen" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AC%E0%A6%82%E0%A6%B6%E0%A6%BE%E0%A6%A3%E0%A7%81" title="বংশাণু – Bangla" lang="bn" hreflang="bn" data-title="বংশাণু" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/%C3%9Bi-tho%C3%A2n-ch%C3%BA" title="Ûi-thoân-chú – Minnan" lang="nan" hreflang="nan" data-title="Ûi-thoân-chú" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Bashkir" lang="ba" hreflang="ba" data-title="Ген" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target"><span>Башҡортса</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Belarusian" lang="be" hreflang="be" data-title="Ген" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Ген" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Bulgarian" lang="bg" hreflang="bg" data-title="Ген" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Gen" title="Gen – Bosnian" lang="bs" hreflang="bs" data-title="Gen" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Gen" title="Gen – Catalan" lang="ca" hreflang="ca" data-title="Gen" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Chuvash" lang="cv" hreflang="cv" data-title="Ген" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target"><span>Чӑвашла</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Gen" title="Gen – Czech" lang="cs" hreflang="cs" data-title="Gen" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-sn mw-list-item"><a href="https://sn.wikipedia.org/wiki/Bari" title="Bari – Shona" lang="sn" hreflang="sn" data-title="Bari" data-language-autonym="ChiShona" data-language-local-name="Shona" class="interlanguage-link-target"><span>ChiShona</span></a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Genyn" title="Genyn – Welsh" lang="cy" hreflang="cy" data-title="Genyn" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target"><span>Cymraeg</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Gen" title="Gen – Danish" lang="da" hreflang="da" data-title="Gen" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-se mw-list-item"><a href="https://se.wikipedia.org/wiki/Gena" title="Gena – Northern Sami" lang="se" hreflang="se" data-title="Gena" data-language-autonym="Davvisámegiella" data-language-local-name="Northern Sami" class="interlanguage-link-target"><span>Davvisámegiella</span></a></li><li class="interlanguage-link interwiki-de badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://de.wikipedia.org/wiki/Gen" title="Gen – German" lang="de" hreflang="de" data-title="Gen" 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/Geen" title="Geen – Estonian" lang="et" hreflang="et" data-title="Geen" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%93%CE%BF%CE%BD%CE%AF%CE%B4%CE%B9%CE%BF" title="Γονίδιο – Greek" lang="el" hreflang="el" data-title="Γονίδιο" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Gen" title="Gen – Spanish" lang="es" hreflang="es" data-title="Gen" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Geno" title="Geno – Esperanto" lang="eo" hreflang="eo" data-title="Geno" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Gene" title="Gene – Basque" lang="eu" hreflang="eu" data-title="Gene" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%98%D9%86" title="ژن – Persian" lang="fa" hreflang="fa" data-title="ژن" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/G%C3%A8ne" title="Gène – French" lang="fr" hreflang="fr" data-title="Gène" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Gen" title="Gen – Western Frisian" lang="fy" hreflang="fy" data-title="Gen" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target"><span>Frysk</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/G%C3%A9in" title="Géin – Irish" lang="ga" hreflang="ga" data-title="Géin" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Xene" title="Xene – Galician" lang="gl" hreflang="gl" data-title="Xene" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-inh mw-list-item"><a href="https://inh.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Ingush" lang="inh" hreflang="inh" data-title="Ген" data-language-autonym="ГӀалгӀай" data-language-local-name="Ingush" class="interlanguage-link-target"><span>ГӀалгӀай</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%9C%A0%EC%A0%84%EC%9E%90" title="유전자 – Korean" lang="ko" hreflang="ko" data-title="유전자" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B3%D5%A5%D5%B6" title="Գեն – Armenian" lang="hy" hreflang="hy" data-title="Գեն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%9C%E0%A5%80%E0%A4%A8" title="जीन – Hindi" lang="hi" hreflang="hi" data-title="जीन" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Gen" title="Gen – Croatian" lang="hr" hreflang="hr" data-title="Gen" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-gor mw-list-item"><a href="https://gor.wikipedia.org/wiki/Gen" title="Gen – Gorontalo" lang="gor" hreflang="gor" data-title="Gen" data-language-autonym="Bahasa Hulontalo" data-language-local-name="Gorontalo" class="interlanguage-link-target"><span>Bahasa Hulontalo</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Geno" title="Geno – Ido" lang="io" hreflang="io" data-title="Geno" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Gen" title="Gen – Indonesian" lang="id" hreflang="id" data-title="Gen" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Gene" title="Gene – Interlingua" lang="ia" hreflang="ia" data-title="Gene" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target"><span>Interlingua</span></a></li><li class="interlanguage-link interwiki-xh mw-list-item"><a href="https://xh.wikipedia.org/wiki/I-gene" title="I-gene – Xhosa" lang="xh" hreflang="xh" data-title="I-gene" data-language-autonym="IsiXhosa" data-language-local-name="Xhosa" class="interlanguage-link-target"><span>IsiXhosa</span></a></li><li class="interlanguage-link interwiki-zu mw-list-item"><a href="https://zu.wikipedia.org/wiki/Isifuzisi" title="Isifuzisi – Zulu" lang="zu" hreflang="zu" data-title="Isifuzisi" data-language-autonym="IsiZulu" data-language-local-name="Zulu" class="interlanguage-link-target"><span>IsiZulu</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Erf%C3%B0av%C3%ADsir" title="Erfðavísir – Icelandic" lang="is" hreflang="is" data-title="Erfðavísir" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Gene" title="Gene – Italian" lang="it" hreflang="it" data-title="Gene" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%92%D7%9F_(%D7%91%D7%99%D7%95%D7%9C%D7%95%D7%92%D7%99%D7%94)" title="גן (ביולוגיה) – Hebrew" lang="he" hreflang="he" data-title="גן (ביולוגיה)" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Gen" title="Gen – Javanese" lang="jv" hreflang="jv" data-title="Gen" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%B5%E0%B2%82%E0%B2%B6%E0%B2%B5%E0%B2%BE%E0%B2%B9%E0%B2%BF" title="ವಂಶವಾಹಿ – Kannada" lang="kn" hreflang="kn" data-title="ವಂಶವಾಹಿ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%92%E1%83%94%E1%83%9C%E1%83%98" title="გენი – Georgian" lang="ka" hreflang="ka" data-title="გენი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Kazakh" lang="kk" hreflang="kk" data-title="Ген" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Jeni" title="Jeni – Swahili" lang="sw" hreflang="sw" data-title="Jeni" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target"><span>Kiswahili</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/J%C3%A8n" title="Jèn – Haitian Creole" lang="ht" hreflang="ht" data-title="Jèn" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-gcr mw-list-item"><a href="https://gcr.wikipedia.org/wiki/J%C3%A8n" title="Jèn – Guianan Creole" lang="gcr" hreflang="gcr" data-title="Jèn" data-language-autonym="Kriyòl gwiyannen" data-language-local-name="Guianan Creole" class="interlanguage-link-target"><span>Kriyòl gwiyannen</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Gen" title="Gen – Kurdish" lang="ku" hreflang="ku" data-title="Gen" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Kyrgyz" lang="ky" hreflang="ky" data-title="Ген" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-lld mw-list-item"><a href="https://lld.wikipedia.org/wiki/Gen" title="Gen – Ladin" lang="lld" hreflang="lld" data-title="Gen" data-language-autonym="Ladin" data-language-local-name="Ladin" class="interlanguage-link-target"><span>Ladin</span></a></li><li class="interlanguage-link interwiki-lo mw-list-item"><a href="https://lo.wikipedia.org/wiki/%E0%BA%A2%E0%BA%B5%E0%BA%99" title="ຢີນ – Lao" lang="lo" hreflang="lo" data-title="ຢີນ" data-language-autonym="ລາວ" data-language-local-name="Lao" class="interlanguage-link-target"><span>ລາວ</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Genum" title="Genum – Latin" lang="la" hreflang="la" data-title="Genum" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/G%C4%93ns" title="Gēns – Latvian" lang="lv" hreflang="lv" data-title="Gēns" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Genas" title="Genas – Lithuanian" lang="lt" hreflang="lt" data-title="Genas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/G%C3%A9n" title="Gén – Hungarian" lang="hu" hreflang="hu" data-title="Gén" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Macedonian" lang="mk" hreflang="mk" data-title="Ген" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%9C%E0%B5%80%E0%B5%BB" title="ജീൻ – Malayalam" lang="ml" hreflang="ml" data-title="ജീൻ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%92%E1%83%94%E1%83%9C%E1%83%98" title="გენი – Mingrelian" lang="xmf" hreflang="xmf" data-title="გენი" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target"><span>მარგალური</span></a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%DA%86%D9%8A%D9%86" title="چين – Egyptian Arabic" lang="arz" hreflang="arz" data-title="چين" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target"><span>مصرى</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Gen" title="Gen – Malay" lang="ms" hreflang="ms" data-title="Gen" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-cdo mw-list-item"><a href="https://cdo.wikipedia.org/wiki/G%C4%AD-%C4%ADng" title="Gĭ-ĭng – Mindong" lang="cdo" hreflang="cdo" data-title="Gĭ-ĭng" data-language-autonym="閩東語 / Mìng-dĕ̤ng-ngṳ̄" data-language-local-name="Mindong" class="interlanguage-link-target"><span>閩東語 / Mìng-dĕ̤ng-ngṳ̄</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Mongolian" lang="mn" hreflang="mn" data-title="Ген" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%99%E1%80%BB%E1%80%AD%E1%80%AF%E1%80%B8%E1%80%9B%E1%80%AD%E1%80%AF%E1%80%B8%E1%80%97%E1%80%AE%E1%80%87" title="မျိုးရိုးဗီဇ – Burmese" lang="my" hreflang="my" data-title="မျိုးရိုးဗီဇ" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target"><span>မြန်မာဘာသာ</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Gen" title="Gen – Dutch" lang="nl" hreflang="nl" data-title="Gen" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ne mw-list-item"><a href="https://ne.wikipedia.org/wiki/%E0%A4%B5%E0%A4%82%E0%A4%B6%E0%A4%BE%E0%A4%A3%E0%A5%81" title="वंशाणु – Nepali" lang="ne" hreflang="ne" data-title="वंशाणु" data-language-autonym="नेपाली" data-language-local-name="Nepali" class="interlanguage-link-target"><span>नेपाली</span></a></li><li class="interlanguage-link interwiki-ja badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://ja.wikipedia.org/wiki/%E9%81%BA%E4%BC%9D%E5%AD%90" title="遺伝子 – Japanese" lang="ja" hreflang="ja" data-title="遺伝子" 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-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/Geen" title="Geen – Northern Frisian" lang="frr" hreflang="frr" data-title="Geen" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target"><span>Nordfriisk</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Gen" title="Gen – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Gen" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Gen" title="Gen – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Gen" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/G%C3%A8n" title="Gèn – Occitan" lang="oc" hreflang="oc" data-title="Gèn" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-om mw-list-item"><a href="https://om.wikipedia.org/wiki/Sanyundee_(gene)" title="Sanyundee (gene) – Oromo" lang="om" hreflang="om" data-title="Sanyundee (gene)" data-language-autonym="Oromoo" data-language-local-name="Oromo" class="interlanguage-link-target"><span>Oromoo</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Gen" title="Gen – Uzbek" lang="uz" hreflang="uz" data-title="Gen" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%9C%E0%A9%80%E0%A8%A8" title="ਜੀਨ – Punjabi" lang="pa" hreflang="pa" data-title="ਜੀਨ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target"><span>ਪੰਜਾਬੀ</span></a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D8%AC%DB%8C%D9%86" title="جین – Western Punjabi" lang="pnb" hreflang="pnb" data-title="جین" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target"><span>پنجابی</span></a></li><li class="interlanguage-link interwiki-blk mw-list-item"><a href="https://blk.wikipedia.org/wiki/%E1%80%85%E1%80%BD%E1%80%AD%E1%80%AF%E1%80%B8%E1%80%81%E1%80%BD%E1%80%AD%E1%80%AF%EA%A9%BB%E1%80%97%E1%80%AE%E1%80%87" title="စွိုးခွိုꩻဗီဇ – Pa'O" lang="blk" hreflang="blk" data-title="စွိုးခွိုꩻဗီဇ" data-language-autonym="ပအိုဝ်ႏဘာႏသာႏ" data-language-local-name="Pa'O" class="interlanguage-link-target"><span>ပအိုဝ်ႏဘာႏသာႏ</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AC%D9%8A%D9%86" title="جين – Pashto" lang="ps" hreflang="ps" data-title="جين" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-jam mw-list-item"><a href="https://jam.wikipedia.org/wiki/Jiin" title="Jiin – Jamaican Creole English" lang="jam" hreflang="jam" data-title="Jiin" data-language-autonym="Patois" data-language-local-name="Jamaican Creole English" class="interlanguage-link-target"><span>Patois</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Gen" title="Gen – Polish" lang="pl" hreflang="pl" data-title="Gen" 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/Gene" title="Gene – Portuguese" lang="pt" hreflang="pt" data-title="Gene" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Gen%C4%83" title="Genă – Romanian" lang="ro" hreflang="ro" data-title="Genă" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-qu mw-list-item"><a href="https://qu.wikipedia.org/wiki/Yawar_saphi" title="Yawar saphi – Quechua" lang="qu" hreflang="qu" data-title="Yawar saphi" data-language-autonym="Runa Simi" data-language-local-name="Quechua" class="interlanguage-link-target"><span>Runa Simi</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Russian" lang="ru" hreflang="ru" data-title="Ген" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sah mw-list-item"><a href="https://sah.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Yakut" lang="sah" hreflang="sah" data-title="Ген" data-language-autonym="Саха тыла" data-language-local-name="Yakut" class="interlanguage-link-target"><span>Саха тыла</span></a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Gene" title="Gene – Scots" lang="sco" hreflang="sco" data-title="Gene" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target"><span>Scots</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Gjeni" title="Gjeni – Albanian" lang="sq" hreflang="sq" data-title="Gjeni" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-scn mw-list-item"><a href="https://scn.wikipedia.org/wiki/Geni" title="Geni – Sicilian" lang="scn" hreflang="scn" data-title="Geni" data-language-autonym="Sicilianu" data-language-local-name="Sicilian" class="interlanguage-link-target"><span>Sicilianu</span></a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%A2%E0%B7%8F%E0%B6%B1" title="ජාන – Sinhala" lang="si" hreflang="si" data-title="ජාන" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Gene" title="Gene – Simple English" lang="en-simple" hreflang="en-simple" data-title="Gene" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sd mw-list-item"><a href="https://sd.wikipedia.org/wiki/%D8%AC%D9%8A%D9%86" title="جين – Sindhi" lang="sd" hreflang="sd" data-title="جين" data-language-autonym="سنڌي" data-language-local-name="Sindhi" class="interlanguage-link-target"><span>سنڌي</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/G%C3%A9n" title="Gén – Slovak" lang="sk" hreflang="sk" data-title="Gén" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Gen" title="Gen – Slovenian" lang="sl" hreflang="sl" data-title="Gen" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-so mw-list-item"><a href="https://so.wikipedia.org/wiki/Hiddoside_(bayoloji)" title="Hiddoside (bayoloji) – Somali" lang="so" hreflang="so" data-title="Hiddoside (bayoloji)" data-language-autonym="Soomaaliga" data-language-local-name="Somali" class="interlanguage-link-target"><span>Soomaaliga</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AC%DB%8C%D9%86" title="جین – Central Kurdish" lang="ckb" hreflang="ckb" data-title="جین" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Serbian" lang="sr" hreflang="sr" data-title="Ген" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Gen" title="Gen – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Gen" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Geeni" title="Geeni – Finnish" lang="fi" hreflang="fi" data-title="Geeni" 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/Gen" title="Gen – Swedish" lang="sv" hreflang="sv" data-title="Gen" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Hene_(biyolohiya)" title="Hene (biyolohiya) – Tagalog" lang="tl" hreflang="tl" data-title="Hene (biyolohiya)" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AE%E0%AE%B0%E0%AE%AA%E0%AE%A3%E0%AF%81" title="மரபணு – Tamil" lang="ta" hreflang="ta" data-title="மரபணு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-te mw-list-item"><a href="https://te.wikipedia.org/wiki/%E0%B0%9C%E0%B0%A8%E0%B1%8D%E0%B0%AF%E0%B1%81%E0%B0%B5%E0%B1%81" title="జన్యువు – Telugu" lang="te" hreflang="te" data-title="జన్యువు" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target"><span>తెలుగు</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A2%E0%B8%B5%E0%B8%99" title="ยีน – Thai" lang="th" hreflang="th" data-title="ยีน" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tg mw-list-item"><a href="https://tg.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Tajik" lang="tg" hreflang="tg" data-title="Ген" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target"><span>Тоҷикӣ</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Gen" title="Gen – Turkish" lang="tr" hreflang="tr" data-title="Gen" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD" title="Ген – Ukrainian" lang="uk" hreflang="uk" data-title="Ген" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D9%88%D8%B1%D8%A7%D8%AB%DB%81" title="وراثہ – Urdu" lang="ur" hreflang="ur" data-title="وراثہ" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-za mw-list-item"><a href="https://za.wikipedia.org/wiki/Gihyinh" title="Gihyinh – Zhuang" lang="za" hreflang="za" data-title="Gihyinh" data-language-autonym="Vahcuengh" data-language-local-name="Zhuang" class="interlanguage-link-target"><span>Vahcuengh</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Gen" title="Gen – Vietnamese" lang="vi" hreflang="vi" data-title="Gen" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh-classical mw-list-item"><a href="https://zh-classical.wikipedia.org/wiki/%E9%81%BA%E5%82%B3%E5%AD%90" title="遺傳子 – Literary Chinese" lang="lzh" hreflang="lzh" data-title="遺傳子" data-language-autonym="文言" data-language-local-name="Literary Chinese" class="interlanguage-link-target"><span>文言</span></a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Hene" title="Hene – Waray" lang="war" hreflang="war" data-title="Hene" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target"><span>Winaray</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E5%9F%BA%E5%9B%A0" title="基因 – Wu" lang="wuu" hreflang="wuu" data-title="基因" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-yi mw-list-item"><a href="https://yi.wikipedia.org/wiki/%D7%92%D7%A2%D7%A0%D7%A2" title="גענע – Yiddish" lang="yi" hreflang="yi" data-title="גענע" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target"><span>ייִדיש</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E5%9F%BA%E5%9B%A0" title="基因 – Cantonese" lang="yue" hreflang="yue" data-title="基因" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li 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vector-pinnable-element"> <div class="vector-pinnable-header vector-appearance-pinnable-header vector-pinnable-header-pinned" data-feature-name="appearance-pinned" data-pinnable-element-id="vector-appearance" data-pinned-container-id="vector-appearance-pinned-container" data-unpinned-container-id="vector-appearance-unpinned-container" > <div class="vector-pinnable-header-label">Appearance</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> <div id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Sequence of DNA that determines traits in an organism</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">This article is about sequences of DNA or RNA that code for functional molecules. For other uses, see <a href="/wiki/Gene_(disambiguation)" class="mw-disambig" title="Gene (disambiguation)">Gene (disambiguation)</a>.</div> <p class="mw-empty-elt"> </p> <table role="presentation" cellpadding="0" style="border-spacing:0; clear:right; float:right;"> <tbody><tr> <td><div class="thumb tright"> <div class="thumbinner" style="width:322px;"> <div class="{{{class}}}" style="position:relative; width:320px; height:185px; overflow:hidden; border:solid #ccc 1px; background-color:var(--background-color-neutral-subtle, #f8f9fa);"> <div style="left:0px; top:0px; width:320px; position:absolute"> <span typeof="mw:File"><a href="/wiki/File:Chromosome_DNA_Gene.svg" title="File:Chromosome DNA Gene.svg"><img alt="A chromosome unravelling into a long string of DNA, a section of which is highlighted as the gene" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/330px-Chromosome_DNA_Gene_unannotated.svg.png" decoding="async" width="320" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/500px-Chromosome_DNA_Gene_unannotated.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/960px-Chromosome_DNA_Gene_unannotated.svg.png 2x" data-file-width="1200" data-file-height="658" /></a></span> </div> <div style="text-align:left; font-size:11px; line-height:110%"> <div style="background-color:transparent; color:var(--color-base, #202122)"><div id="annotation_137x52" style="position:absolute; left:137px; top:52px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Chromosome" title="Chromosome">Chromosome </a> </span></div> <div id="annotation_210x52" style="position:absolute; left:210px; top:52px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> (<a href="/wiki/Base_pair#Length_measurements" title="Base pair">10<sup>7</sup> - 10<sup>10</sup> bp</a>)</span></div> <div id="annotation_25x107" style="position:absolute; left:25px; top:107px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/DNA" title="DNA">DNA </a> </span></div> <div id="annotation_137x118" style="position:absolute; left:137px; top:118px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a class="mw-selflink selflink">Gene </a> </span></div> <div id="annotation_173x118" style="position:absolute; left:173px; top:118px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> (<a href="/wiki/Base_pair#Length_measurements" title="Base pair">10<sup>3</sup> - 10<sup>6</sup> bp </a>)</span></div> <div id="annotation_165x165" style="position:absolute; left:165px; top:165px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Phenotype" title="Phenotype">Function </a> </span></div></div> </div> </div> <div class="thumbcaption" style="clear:left"><div style="float:left;margin-right:0.5em"><span typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/20px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/40px-Interactive_icon.svg.png 1.5x" data-file-width="133" data-file-height="200" /></span></span></div>A chromosome and its <a href="/wiki/Nuclear_organization" title="Nuclear organization">packaged</a> long strand of DNA unraveled. The DNA's <a href="/wiki/Base_pair" title="Base pair">base pairs</a> encode genes, which provide functions. A human DNA can have up to 500 million base pairs with thousands of genes.</div> </div> </div> </td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · 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.sidebar:not(.notheme) .sidebar-list-title,html.skin-theme-clientpref-night .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-list-title,html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media print{body.ns-0 .mw-parser-output .sidebar{display:none!important}}</style><style data-mw-deduplicate="TemplateStyles:r1066933788">.mw-parser-output .excerpt-hat .mw-editsection-like{font-style:normal}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788" /><table class="sidebar sidebar-collapse nomobile nowraplinks" style="width:17em;border:2px solid light-dark(#90db90,#024c02)"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:175%;font-weight:bold"><a href="/wiki/Genetics" title="Genetics">Genetics</a></th></tr><tr><td class="sidebar-image"><span typeof="mw:File"><a href="/wiki/File:Hybridogenesis_in_water_frogs_gametes.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/150px-Hybridogenesis_in_water_frogs_gametes.svg.png" decoding="async" width="150" height="74" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/225px-Hybridogenesis_in_water_frogs_gametes.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/300px-Hybridogenesis_in_water_frogs_gametes.svg.png 2x" data-file-width="225" data-file-height="111" /></a></span></td></tr><tr><td class="sidebar-content" style="background:light-dark(#e7f4e7,#243124);color:inherit;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:100%;font-weight:bold;;color: var(--color-base)">Key components</div><div class="sidebar-list-content mw-collapsible-content hlist"><div class="excerpt-block"><div class="excerpt"> <ul><li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotide</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul></div></div> <hr /> <ul><li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:light-dark(#e7f4e7,#243124);color:inherit;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:100%;font-weight:bold;;color: var(--color-base)">History and topics</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Evolution" title="Evolution">Evolution</a> (<a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular</a>)</li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular genetics</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:light-dark(#e7f4e7,#243124);color:inherit;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:100%;font-weight:bold;;color: var(--color-base)">Research</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><span class="nowrap"><a href="/wiki/Genomics" title="Genomics">Genomics</a> (<span class="noviewer" typeof="mw:File"><span title="Template"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Symbol_template_class_pink.svg/20px-Symbol_template_class_pink.svg.png" decoding="async" width="12" height="12" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Symbol_template_class_pink.svg/40px-Symbol_template_class_pink.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Template:Genomics" title="Template:Genomics">template</a>)</span></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</a></li></ul> <hr /> <ul><li><a href="/wiki/Outline_of_genetics#Branches_of_genetics" title="Outline of genetics">Branches of genetics</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:light-dark(#e7f4e7,#243124);color:inherit;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields</div><div class="sidebar-list-content mw-collapsible-content hlist"><div class="excerpt-block"><div class="excerpt"> <ul><li><a href="/wiki/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul></div></div></div></div></td> </tr><tr><td class="sidebar-content" style="background:light-dark(#e7f4e7,#243124);color:inherit;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:light-dark(#90db90,#024c02) !important;color:inherit;font-size:100%;font-weight:bold;;color: var(--color-base)">Personalized medicine</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Personalized_medicine" title="Personalized medicine">Personalized medicine</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Genetics" title="Category:Genetics">Category</a></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genetics_sidebar" title="Template:Genetics sidebar"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetics_sidebar" title="Template talk:Genetics sidebar"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetics_sidebar" title="Special:EditPage/Template:Genetics sidebar"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p>In <a href="/wiki/Biology" title="Biology">biology</a>, the word <b>gene</b> has two meanings. The Mendelian gene is a basic unit of <a href="/wiki/Heredity" title="Heredity">heredity</a>. The molecular gene is a sequence of <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a> in <a href="/wiki/DNA" title="DNA">DNA</a> that is transcribed to produce a functional <a href="/wiki/RNA" title="RNA">RNA</a>. There are two types of molecular genes: protein-coding genes and non-coding genes.<sup id="cite_ref-mendgene2_1-0" class="reference"><a href="#cite_note-mendgene2-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MedlinePlus_2020_2-0" class="reference"><a href="#cite_note-MedlinePlus_2020-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hirsch_2002_p._3-0" class="reference"><a href="#cite_note-Hirsch_2002_p.-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> During <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> (the synthesis of <a href="/wiki/Gene_product" title="Gene product">RNA or protein</a> from a gene), DNA is first <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">copied into RNA</a>. RNA can be <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">directly functional</a> or be the intermediate <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">template</a> for the synthesis of a protein. </p><p>The transmission of genes to an organism's <a href="/wiki/Offspring" title="Offspring">offspring</a>, is the basis of the inheritance of <a href="/wiki/Phenotypic_trait" title="Phenotypic trait">phenotypic traits</a> from one generation to the next. These genes make up different DNA sequences, together called a <a href="/wiki/Genotype" title="Genotype">genotype</a>, that is specific to every given individual, within the <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a> of the <a href="/wiki/Population_(biology)" class="mw-redirect" title="Population (biology)">population</a> of a given <a href="/wiki/Species" title="Species">species</a>. The genotype, along with environmental and developmental factors, ultimately determines the <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> of the individual. </p><p>Most biological traits occur under the combined influence of <a href="/wiki/Polygene" title="Polygene">polygenes</a> (a set of different genes) and <a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">gene–environment interactions</a>. Some genetic traits are instantly visible, such as <a href="/wiki/Eye_color" title="Eye color">eye color</a> or the number of limbs, others are not, such as <a href="/wiki/Blood_type" title="Blood type">blood type</a>, the risk for specific diseases, or the thousands of basic <a href="/wiki/Biochemistry" title="Biochemistry">biochemical</a> processes that constitute <a href="/wiki/Life" title="Life">life</a>. A gene can acquire <a href="/wiki/Mutation" title="Mutation">mutations</a> in its <a href="/wiki/Gene_sequence" class="mw-redirect" title="Gene sequence">sequence</a>, leading to different variants, known as <a href="/wiki/Allele" title="Allele">alleles</a>, in the <a href="/wiki/Population" title="Population">population</a>. These alleles encode slightly different versions of a gene, which may cause different <a href="/wiki/Phenotypical" class="mw-redirect" title="Phenotypical">phenotypical</a> traits.<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> Genes <a href="/wiki/Evolution" title="Evolution">evolve</a> due to <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> or <a href="/wiki/Survival_of_the_fittest" title="Survival of the fittest">survival of the fittest</a> and <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a> of the alleles. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Definitions">Definitions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=1" title="Edit section: Definitions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are many different ways to use the term "gene" based on different aspects of their inheritance, selection, biological function, or molecular structure but most of these definitions fall into two categories, the Mendelian gene or the molecular gene.<sup id="cite_ref-mendgene2_1-1" class="reference"><a href="#cite_note-mendgene2-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kampourakis_5-0" class="reference"><a href="#cite_note-Kampourakis-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Gericke_6-0" class="reference"><a href="#cite_note-Gericke-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><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> </p><p>The Mendelian gene is the classical gene of genetics and it refers to any heritable trait. This is the gene described in <i>The Selfish Gene</i>.<sup id="cite_ref-simply2_9-0" class="reference"><a href="#cite_note-simply2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> More thorough discussions of this version of a gene can be found in the articles <i><a href="/wiki/Genetics" title="Genetics">Genetics</a></i> and <i><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></i>. </p><p>The molecular gene definition is more commonly used across biochemistry, molecular biology, and most of genetics—the gene that is described in terms of DNA sequence.<sup id="cite_ref-mendgene2_1-2" class="reference"><a href="#cite_note-mendgene2-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> There are many different definitions of this gene—some of which are misleading or incorrect.<sup id="cite_ref-Kampourakis_5-1" class="reference"><a href="#cite_note-Kampourakis-5"><span class="cite-bracket">[</span>5<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>Very early work in the field that became <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a> suggested the concept that <a href="/wiki/One_gene%E2%80%93one_enzyme_hypothesis" title="One gene–one enzyme hypothesis">one gene makes one protein</a> (originally 'one gene – one enzyme').<sup id="cite_ref-pmid16588492_11-0" class="reference"><a href="#cite_note-pmid16588492-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid15020400_12-0" class="reference"><a href="#cite_note-pmid15020400-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> However, genes that produce repressor RNAs were proposed in the 1950s<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and by the 1960s, textbooks were using molecular gene definitions that included those that specified functional RNA molecules such as ribosomal RNA and tRNA (noncoding genes) as well as protein-coding genes.<sup id="cite_ref-Watson_14-0" class="reference"><a href="#cite_note-Watson-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>This idea of two kinds of genes is still part of the definition of a gene in most textbooks. For example, </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>The primary function of the genome is to produce RNA molecules. Selected portions of the DNA nucleotide sequence are copied into a corresponding RNA nucleotide sequence, which either encodes a protein (if it is an mRNA) or forms a 'structural' RNA, such as a transfer RNA (tRNA) or ribosomal RNA (rRNA) molecule. Each region of the DNA helix that produces a functional RNA molecule constitutes a gene.<sup id="cite_ref-Alberts_15-0" class="reference"><a href="#cite_note-Alberts-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup></p></blockquote> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712" /><blockquote class="templatequote"><p>We define a gene as a DNA sequence that is transcribed. This definition includes genes that do not encode proteins (not all transcripts are messenger RNA). The definition normally excludes regions of the genome that control transcription but are not themselves transcribed. We will encounter some exceptions to our definition of a gene - surprisingly, there is no definition that is entirely satisfactory.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></p></blockquote> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712" /><blockquote class="templatequote"><p>A gene is a DNA sequence that codes for a diffusible product. This product may be protein (as is the case in the majority of genes) or may be RNA (as is the case of genes that code for tRNA and rRNA). The crucial feature is that the product diffuses away from its site of synthesis to act elsewhere.<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></p></blockquote> <p>The important parts of such definitions are: (1) that a gene corresponds to a transcription unit; (2) that genes produce both mRNA and noncoding RNAs; and (3) regulatory sequences control gene expression but are not part of the gene itself. However, there is one other important part of the definition and it is emphasized in Kostas Kampourakis' book <i>Making Sense of Genes</i>. </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712" /><blockquote class="templatequote"><p>Therefore in this book I will consider genes as DNA sequences encoding information for functional products, be it proteins or RNA molecules. With 'encoding information', I mean that the DNA sequence is used as a template for the production of an RNA molecule or a protein that performs some function.<sup id="cite_ref-Kampourakis_5-2" class="reference"><a href="#cite_note-Kampourakis-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup></p></blockquote> <p>The emphasis on function is essential because there are stretches of DNA that produce non-functional transcripts and they do not qualify as genes. These include obvious examples such as transcribed pseudogenes as well as less obvious examples such as junk RNA produced as noise due to transcription errors. In order to qualify as a true gene, by this definition, one has to prove that the transcript has a biological function.<sup id="cite_ref-Kampourakis_5-3" class="reference"><a href="#cite_note-Kampourakis-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Early speculations on the size of a typical gene were based on high-resolution genetic mapping and on the size of proteins and RNA molecules. A length of 1500 base pairs seemed reasonable at the time (1965).<sup id="cite_ref-Watson_14-1" class="reference"><a href="#cite_note-Watson-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> This was based on the idea that the gene was the DNA that was directly responsible for production of the functional product. The discovery of introns in the 1970s meant that many eukaryotic genes were much larger than the size of the functional product would imply. Typical mammalian protein-coding genes, for example, are about 62,000 base pairs in length (transcribed region) and since there are about 20,000 of them they occupy about 35–40% of the mammalian genome (including the human genome).<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><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> </p><p>In spite of the fact that both protein-coding genes and noncoding genes have been known for more than 50 years, there are still a number of textbooks, websites, and scientific publications that define a gene as a DNA sequence that specifies a protein. In other words, the definition is restricted to protein-coding genes. Here is an example from a recent article in American Scientist. </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712" /><blockquote class="templatequote"><p>... to truly assess the potential significance of de novo genes, we relied on a strict definition of the word "gene" with which nearly every expert can agree. First, in order for a nucleotide sequence to be considered a true gene, an open reading frame (ORF) must be present. The ORF can be thought of as the "gene itself"; it begins with a starting mark common for every gene and ends with one of three possible finish line signals. One of the key enzymes in this process, the RNA polymerase, zips along the strand of DNA like a train on a monorail, transcribing it into its messenger RNA form. This point brings us to our second important criterion: A true gene is one that is both transcribed and translated. That is, a true gene is first used as a template to make transient messenger RNA, which is then translated into a protein.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></p></blockquote> <p>This restricted definition is so common that it has spawned many recent articles that criticize this "standard definition" and call for a new expanded definition that includes noncoding genes. However, some modern writers still do not acknowledge noncoding genes although this so-called "new" definition has been recognised for more than half a century.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Although some definitions can be more broadly applicable than others, the fundamental complexity of biology means that no definition of a gene can capture all aspects perfectly. Not all genomes are DNA (e.g. <a href="/wiki/RNA_virus" title="RNA virus">RNA viruses</a>),<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> bacterial <a href="/wiki/Operon" title="Operon">operons</a> are multiple protein-coding regions transcribed into single large mRNAs, <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a> enables a single genomic region to encode multiple district products and <a href="/wiki/Trans-splicing" title="Trans-splicing">trans-splicing</a> concatenates mRNAs from shorter coding sequence across the genome.<sup id="cite_ref-Rethink_26-0" class="reference"><a href="#cite_note-Rethink-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Since molecular definitions exclude elements such as introns, promotors, and other <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">regulatory regions</a>, these are instead thought of as "associated" with the gene and affect its function. </p><p>An even broader operational definition is sometimes used to encompass the complexity of these diverse phenomena, where a gene is defined as a union of genomic sequences encoding a coherent set of potentially overlapping functional products.<sup id="cite_ref-Gerstein_29-0" class="reference"><a href="#cite_note-Gerstein-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This definition categorizes genes by their functional products (proteins or RNA) rather than their specific DNA loci, with regulatory elements classified as <i>gene-associated</i> regions.<sup id="cite_ref-Gerstein_29-1" class="reference"><a href="#cite_note-Gerstein-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=2" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/History_of_genetics" title="History of genetics">History of genetics</a></div> <div class="mw-heading mw-heading3"><h3 id="Discovery_of_discrete_inherited_units">Discovery of discrete inherited units</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=3" title="Edit section: Discovery of discrete inherited units"><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:Gregor_Mendel_-_American_Breeders_Magazine,_1910.jpg" class="mw-file-description"><img alt="Photograph of Gregor Mendel" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg/180px-Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg" decoding="async" width="180" height="231" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg/270px-Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg/360px-Gregor_Mendel_-_American_Breeders_Magazine%2C_1910.jpg 2x" data-file-width="780" data-file-height="1000" /></a><figcaption>Gregor Mendel</figcaption></figure> <p>The existence of discrete inheritable units was first suggested by <a href="/wiki/Gregor_Mendel" title="Gregor Mendel">Gregor Mendel</a> (1822–1884).<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> From 1857 to 1864, in <a href="/wiki/Brno" title="Brno">Brno</a>, <a href="/wiki/Austrian_Empire" title="Austrian Empire">Austrian Empire</a> (today's Czech Republic), he studied inheritance patterns in 8000 common edible <a href="/wiki/Pea" title="Pea">pea plants</a>, tracking distinct traits from parent to offspring. He described these mathematically as 2<sup>n</sup> combinations where n is the number of differing characteristics in the original peas. Although he did not use the term <i>gene</i>, he explained his results in terms of discrete inherited units that give rise to observable physical characteristics. This description prefigured <a href="/wiki/Wilhelm_Johannsen" title="Wilhelm Johannsen">Wilhelm Johannsen</a>'s distinction between <a href="/wiki/Genotype" title="Genotype">genotype</a> (the genetic material of an organism) and <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> (the observable traits of that organism). Mendel was also the first to demonstrate <a href="/wiki/Independent_assortment" class="mw-redirect" title="Independent assortment">independent assortment</a>, the distinction between <a href="/wiki/Dominant_gene" class="mw-redirect" title="Dominant gene">dominant</a> and <a href="/wiki/Recessive" class="mw-redirect" title="Recessive">recessive</a> traits, the distinction between a <a href="/wiki/Heterozygote" class="mw-redirect" title="Heterozygote">heterozygote</a> and <a href="/wiki/Homozygote" class="mw-redirect" title="Homozygote">homozygote</a>, and the phenomenon of discontinuous inheritance. </p><p>Prior to Mendel's work, the dominant theory of heredity was one of <a href="/wiki/Blending_inheritance" title="Blending inheritance">blending inheritance</a>,<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> which suggested that each parent contributed fluids to the fertilization process and that the traits of the parents blended and mixed to produce the offspring. <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> developed a theory of inheritance he termed <a href="/wiki/Pangenesis" title="Pangenesis">pangenesis</a>, from <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> pan ("all, whole") and genesis ("birth") / genos ("origin").<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Darwin used the term <i><a href="/wiki/Gemmule_(pangenesis)" class="mw-redirect" title="Gemmule (pangenesis)">gemmule</a></i> to describe hypothetical particles that would mix during reproduction. </p><p>Mendel's work went largely unnoticed after its first publication in 1866, but was rediscovered in the late 19th century by <a href="/wiki/Hugo_de_Vries" title="Hugo de Vries">Hugo de Vries</a>, <a href="/wiki/Carl_Correns" title="Carl Correns">Carl Correns</a>, and <a href="/wiki/Erich_von_Tschermak" title="Erich von Tschermak">Erich von Tschermak</a>, who (claimed to have) reached similar conclusions in their own research.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Specifically, in 1889, Hugo de Vries published his book <i>Intracellular Pangenesis</i>,<sup id="cite_ref-de_Vries_1889_35-0" class="reference"><a href="#cite_note-de_Vries_1889-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> in which he postulated that different characters have individual hereditary carriers and that inheritance of specific traits in organisms comes in particles. De Vries called these units "pangenes" (<i>Pangens</i> in German), after Darwin's 1868 pangenesis theory. </p><p>Twenty years later, in 1909, <a href="/wiki/Wilhelm_Johannsen" title="Wilhelm Johannsen">Wilhelm Johannsen</a> introduced the term "gene" (inspired by the <a href="/wiki/Ancient_Greek" title="Ancient Greek">ancient Greek</a>: γόνος, <i>gonos</i>, meaning offspring and procreation)<sup id="cite_ref-Johannsen_36-0" class="reference"><a href="#cite_note-Johannsen-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> and, in 1906, <a href="/wiki/William_Bateson" title="William Bateson">William Bateson</a>, that of "<a href="/wiki/Genetics" title="Genetics">genetics</a>"<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><sup id="cite_ref-Gerstein_29-2" class="reference"><a href="#cite_note-Gerstein-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> while <a href="/wiki/Eduard_Strasburger" title="Eduard Strasburger">Eduard Strasburger</a>, among others, still used the term "pangene" for the fundamental physical and functional unit of heredity.<sup id="cite_ref-de_Vries_1889_35-1" class="reference"><a href="#cite_note-de_Vries_1889-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: Translator's preface, viii">: Translator's preface, viii </span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Discovery_of_DNA">Discovery of DNA</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=4" title="Edit section: Discovery of DNA"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Advances in understanding genes and inheritance continued throughout the 20th century. <a href="/wiki/Deoxyribonucleic_acid" class="mw-redirect" title="Deoxyribonucleic acid">Deoxyribonucleic acid</a> (DNA) was shown to be the molecular repository of genetic information by experiments in the 1940s to 1950s.<sup id="cite_ref-Avery_et_al_38-0" class="reference"><a href="#cite_note-Avery_et_al-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> The structure of DNA was studied by <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a> and <a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Maurice Wilkins</a> using <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>, which led <a href="/wiki/James_D._Watson" class="mw-redirect" title="James D. Watson">James D. Watson</a> and <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> to publish a model of the double-stranded DNA molecule whose paired <a href="/wiki/Nucleotide_base" title="Nucleotide base">nucleotide bases</a> indicated a compelling hypothesis for the mechanism of genetic replication.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-watsoncrick_1953a_41-0" class="reference"><a href="#cite_note-watsoncrick_1953a-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p><p>In the early 1950s the prevailing view was that the genes in a chromosome acted like discrete entities arranged like beads on a string. The experiments of <a href="/wiki/Seymour_Benzer" title="Seymour Benzer">Benzer</a> using <a href="/wiki/Mutant" title="Mutant">mutants</a> defective in the <a href="/wiki/T4_rII_system" title="T4 rII system">rII region of bacteriophage T4</a> (1955–1959) showed that individual genes have a simple linear structure and are likely to be equivalent to a linear section of DNA.<sup id="cite_ref-pmid16589677_42-0" class="reference"><a href="#cite_note-pmid16589677-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid16590553_43-0" class="reference"><a href="#cite_note-pmid16590553-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>Collectively, this body of research established the <a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">central dogma of molecular biology</a>, which states that <a href="/wiki/Protein" title="Protein">proteins</a> are translated from <a href="/wiki/RNA" title="RNA">RNA</a>, which is transcribed from <a href="/wiki/DNA" title="DNA">DNA</a>. This dogma has since been shown to have exceptions, such as <a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">reverse transcription</a> in <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>. The modern study of <a href="/wiki/Genetics" title="Genetics">genetics</a> at the level of DNA is known as <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a>. </p><p>In 1972, <a href="/wiki/Walter_Fiers" title="Walter Fiers">Walter Fiers</a> and his team were the first to determine the sequence of a gene: that of <a href="/wiki/Bacteriophage_MS2" title="Bacteriophage MS2">bacteriophage MS2</a> coat protein.<sup id="cite_ref-Min_1972_44-0" class="reference"><a href="#cite_note-Min_1972-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> The subsequent development of <a href="/wiki/Sanger_sequencing" title="Sanger sequencing">chain-termination</a> <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> in 1977 by <a href="/wiki/Frederick_Sanger" title="Frederick Sanger">Frederick Sanger</a> improved the efficiency of sequencing and turned it into a routine laboratory tool.<sup id="cite_ref-sanger_et_al_45-0" class="reference"><a href="#cite_note-sanger_et_al-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> An automated version of the Sanger method was used in early phases of the <a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a>.<sup id="cite_ref-scitable_adams_46-0" class="reference"><a href="#cite_note-scitable_adams-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Modern_synthesis_and_its_successors">Modern synthesis and its successors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=5" title="Edit section: Modern synthesis and its successors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis (20th century)</a></div> <p>The theories developed in the early 20th century to integrate <a href="/wiki/Mendelian_genetics" class="mw-redirect" title="Mendelian genetics">Mendelian genetics</a> with <a href="/wiki/Darwinian_evolution" class="mw-redirect" title="Darwinian evolution">Darwinian evolution</a> are called the <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern synthesis</a>, a term introduced by <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a>.<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>This view of evolution was emphasized by <a href="/wiki/George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">George C. Williams</a>' <a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">gene-centric view of evolution</a>. He proposed that the Mendelian gene is a <a href="/wiki/Unit_of_selection" title="Unit of selection">unit</a> of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> with the definition: "that which segregates and recombines with appreciable frequency."<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><sup class="reference nowrap"><span title="Page / location: 24">: 24 </span></sup> Related ideas emphasizing the centrality of Mendelian genes and the importance of natural selection in evolution were popularized by <a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a>.<sup id="cite_ref-simply2_9-1" class="reference"><a href="#cite_note-simply2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p><p>The development of the <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of evolution</a> in the late 1960s led to the recognition that random genetic drift is a major player in evolution and that neutral theory should be the null hypothesis of molecular evolution.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> This led to the construction of <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic trees</a> and the development of the <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a>, which is the basis of all dating techniques using DNA sequences. These techniques are not confined to molecular gene sequences but can be used on all DNA segments in the genome. </p> <div class="mw-heading mw-heading2"><h2 id="Molecular_basis">Molecular basis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=6" title="Edit section: Molecular basis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/DNA" title="DNA">DNA</a></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_chemical_structure_2.svg" class="mw-file-description"><img alt="DNA chemical structure diagram showing how the double helix consists of two chains of sugar-phosphate backbone with bases pointing inward and specifically base pairing A to T and C to G with hydrogen bonds." src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/DNA_chemical_structure_2.svg/330px-DNA_chemical_structure_2.svg.png" decoding="async" width="330" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/DNA_chemical_structure_2.svg/495px-DNA_chemical_structure_2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b2/DNA_chemical_structure_2.svg/660px-DNA_chemical_structure_2.svg.png 2x" data-file-width="1200" data-file-height="532" /></a><figcaption>The chemical structure of a four base pair fragment of a <a href="/wiki/DNA" title="DNA">DNA</a> <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a>. The <a href="/wiki/Deoxyribose" title="Deoxyribose">sugar</a>-<a href="/wiki/Phosphate" title="Phosphate">phosphate</a> backbone chains run in opposite directions with the <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">bases</a> pointing inward, <a href="/wiki/Base-pair" class="mw-redirect" title="Base-pair">base-pairing</a> <a href="/wiki/Adenine" title="Adenine">A</a> to <a href="/wiki/Thymine" title="Thymine">T</a> and <a href="/wiki/Cytosine" title="Cytosine">C</a> to <a href="/wiki/Guanine" title="Guanine">G</a> with <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a>. </figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="DNA">DNA</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=7" title="Edit section: DNA"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The vast majority of organisms encode their genes in long strands of <a href="/wiki/DNA" title="DNA">DNA</a> (deoxyribonucleic acid). DNA consists of a <a href="/wiki/Polymer" title="Polymer">chain</a> made from four types of <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> subunits, each composed of: a five-carbon sugar (<a href="/wiki/Deoxyribose" title="Deoxyribose">2-deoxyribose</a>), a <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> group, and one of the four <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">bases</a> <a href="/wiki/Adenine" title="Adenine">adenine</a>, <a href="/wiki/Cytosine" title="Cytosine">cytosine</a>, <a href="/wiki/Guanine" title="Guanine">guanine</a>, and <a href="/wiki/Thymine" title="Thymine">thymine</a>.<sup id="cite_ref-MBOC_51-0" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 2.1">: 2.1 </span></sup> </p><p>Two chains of DNA twist around each other to form a DNA <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a> with the phosphate–sugar backbone spiralling around the outside, and the bases pointing inward with adenine <a href="/wiki/Base_pair" title="Base pair">base pairing</a> to thymine and guanine to cytosine. The specificity of base pairing occurs because adenine and thymine align to form two <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a>, whereas cytosine and guanine form three hydrogen bonds. The two strands in a double helix must, therefore, be <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a>, with their sequence of bases matching such that the adenines of one strand are paired with the thymines of the other strand, and so on.<sup id="cite_ref-MBOC_51-1" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 4.1">: 4.1 </span></sup> </p><p>Due to the chemical composition of the <a href="/wiki/Pentose" title="Pentose">pentose</a> residues of the bases, DNA strands have directionality. One end of a DNA <a href="/wiki/Polymer" title="Polymer">polymer</a> contains an exposed <a href="/wiki/Hydroxyl" class="mw-redirect" title="Hydroxyl">hydroxyl</a> group on the <a href="/wiki/Deoxyribose" title="Deoxyribose">deoxyribose</a>; this is known as the <a href="/wiki/3%27_end" class="mw-redirect" title="3' end">3' end</a> of the molecule. The other end contains an exposed <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> group; this is the <a href="/wiki/5%27_end" class="mw-redirect" title="5' end">5' end</a>. The two strands of a double-helix run in opposite directions. Nucleic acid synthesis, including <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> and <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> occurs in the 5'→3' direction, because new nucleotides are added via a <a href="/wiki/Dehydration_reaction" title="Dehydration reaction">dehydration reaction</a> that uses the exposed 3' hydroxyl as a <a href="/wiki/Nucleophile" title="Nucleophile">nucleophile</a>.<sup id="cite_ref-Stryer_2002_52-0" class="reference"><a href="#cite_note-Stryer_2002-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 27.2">: 27.2 </span></sup> </p><p>The <a href="/wiki/Gene_expression" title="Gene expression">expression</a> of genes encoded in DNA begins by <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribing</a> the gene into <a href="/wiki/RNA" title="RNA">RNA</a>, a second type of nucleic acid that is very similar to DNA, but whose monomers contain the sugar <a href="/wiki/Ribose" title="Ribose">ribose</a> rather than <a href="/wiki/Deoxyribose" title="Deoxyribose">deoxyribose</a>. RNA also contains the base <a href="/wiki/Uracil" title="Uracil">uracil</a> in place of <a href="/wiki/Thymine" title="Thymine">thymine</a>. RNA molecules are less stable than DNA and are typically single-stranded. Genes that encode proteins are composed of a series of three-<a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> sequences called <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>, which serve as the "words" in the genetic "language". The <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> specifies the correspondence during <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">protein translation</a> between codons and <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a>. The genetic code is nearly the same for all known organisms.<sup id="cite_ref-MBOC_51-2" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 4.1">: 4.1 </span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Chromosomes">Chromosomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=8" title="Edit section: Chromosomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:NHGRI_human_male_karyotype.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/53/NHGRI_human_male_karyotype.png/250px-NHGRI_human_male_karyotype.png" decoding="async" width="220" height="172" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/53/NHGRI_human_male_karyotype.png/330px-NHGRI_human_male_karyotype.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/53/NHGRI_human_male_karyotype.png/500px-NHGRI_human_male_karyotype.png 2x" data-file-width="3084" data-file-height="2417" /></a><figcaption><a href="/wiki/Micrograph" title="Micrograph">Micrographic</a> <a href="/wiki/Karyotype" title="Karyotype">karyogram</a> of human male, showing 23 pairs of chromosomes. The largest <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> are around 10 times the size of the smallest.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_karyotype_with_bands_and_sub-bands.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/250px-Human_karyotype_with_bands_and_sub-bands.png" decoding="async" width="220" height="364" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/330px-Human_karyotype_with_bands_and_sub-bands.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/500px-Human_karyotype_with_bands_and_sub-bands.png 2x" data-file-width="9684" data-file-height="16008" /></a><figcaption>Schematic <a href="/wiki/Karyotype" title="Karyotype">karyogram</a> of a human, with annotated <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">bands and sub-bands</a>. It shows dark and white regions on <a href="/wiki/G_banding" title="G banding">G banding</a>. It shows 22 <a href="/wiki/Homologous_chromosome" title="Homologous chromosome">homologous chromosomes</a>, both the male (XY) and female (XX) versions of the <a href="/wiki/Sex_chromosome" title="Sex chromosome">sex chromosome</a> (bottom right), as well as the <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">mitochondrial genome</a> (at bottom left). <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Karyotype" title="Karyotype">Karyotype</a></div></figcaption></figure> <p>The total complement of genes in an organism or cell is known as its <a href="/wiki/Genome" title="Genome">genome</a>, which may be stored on one or more <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>. A chromosome consists of a single, very long DNA helix on which thousands of genes are encoded.<sup id="cite_ref-MBOC_51-3" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 4.2">: 4.2 </span></sup> The region of the chromosome at which a particular gene is located is called its <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">locus</a>. Each locus contains one <a href="/wiki/Allele" title="Allele">allele</a> of a gene; however, members of a population may have different alleles at the locus, each with a slightly different gene sequence. </p><p>The majority of <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> genes are stored on a set of large, linear chromosomes. The chromosomes are packed within the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> in complex with storage proteins called <a href="/wiki/Histone" title="Histone">histones</a> to form a unit called a <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a>. DNA packaged and condensed in this way is called <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>.<sup id="cite_ref-MBOC_51-4" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 4.2">: 4.2 </span></sup> The manner in which DNA is stored on the histones, as well as chemical modifications of the histone itself, regulate whether a particular region of DNA is accessible for <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>. In addition to genes, eukaryotic chromosomes contain sequences involved in ensuring that the DNA is copied without degradation of end regions and sorted into daughter cells during cell division: <a href="/wiki/Replication_origin" class="mw-redirect" title="Replication origin">replication origins</a>, <a href="/wiki/Telomere" title="Telomere">telomeres</a>, and the <a href="/wiki/Centromere" title="Centromere">centromere</a>.<sup id="cite_ref-MBOC_51-5" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 4.2">: 4.2 </span></sup> Replication origins are the sequence regions where <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> is initiated to make two copies of the chromosome. Telomeres are long stretches of repetitive sequences that cap the ends of the linear chromosomes and prevent degradation of coding and regulatory regions during <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. The length of the telomeres decreases each time the genome is replicated and has been implicated in the <a href="/wiki/Aging" class="mw-redirect" title="Aging">aging</a> process.<sup id="cite_ref-Braig_54-0" class="reference"><a href="#cite_note-Braig-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> The centromere is required for binding <a href="/wiki/Spindle_fibre" class="mw-redirect" title="Spindle fibre">spindle fibres</a> to separate sister chromatids into daughter cells during <a href="/wiki/Cell_division" title="Cell division">cell division</a>.<sup id="cite_ref-MBOC_51-6" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 18.2">: 18.2 </span></sup> </p><p><a href="/wiki/Prokaryote" title="Prokaryote">Prokaryotes</a> (<a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>) typically store their genomes on a single, large, <a href="/wiki/Circular_chromosome" title="Circular chromosome">circular chromosome</a>. Similarly, some eukaryotic <a href="/wiki/Organelle" title="Organelle">organelles</a> contain a remnant circular chromosome with a small number of genes.<sup id="cite_ref-MBOC_51-7" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 14.4">: 14.4 </span></sup> Prokaryotes sometimes supplement their chromosome with additional small circles of DNA called <a href="/wiki/Plasmid" title="Plasmid">plasmids</a>, which usually encode only a few genes and are transferable between individuals. For example, the genes for <a href="/wiki/Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a> are usually encoded on bacterial plasmids and can be passed between individual cells, even those of different species, via <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>.<sup id="cite_ref-bennett_55-0" class="reference"><a href="#cite_note-bennett-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>Whereas the chromosomes of prokaryotes are relatively gene-dense, those of eukaryotes often contain regions of DNA that serve no obvious function. Simple single-celled eukaryotes have relatively small amounts of such DNA, whereas the genomes of complex <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellular organisms</a>, including humans, contain an absolute majority of DNA without an identified function.<sup id="cite_ref-IHSGC2004_56-0" class="reference"><a href="#cite_note-IHSGC2004-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> This DNA has often been referred to as "<a href="/wiki/Junk_DNA" title="Junk DNA">junk DNA</a>". However, more recent analyses suggest that, although protein-coding DNA makes up barely 2% of the <a href="/wiki/Human_genome" title="Human genome">human genome</a>, about 80% of the bases in the genome may be expressed, so the term "junk DNA" may be a misnomer.<sup id="cite_ref-Rethink_26-1" class="reference"><a href="#cite_note-Rethink-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structure_and_function">Structure and function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=9" title="Edit section: Structure and function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Structure">Structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=10" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table style="clear:right; float: right;"> <tbody><tr> <td> <table role="presentation" cellpadding="0" style="border-spacing:0; clear:right; float:right;"> <tbody><tr> <td><div class="thumb tright"> <div class="thumbinner" style="width:602px;"> <div class="skin-invert" style="position:relative; width:600px; height:325px; overflow:hidden; border:solid #ccc 1px; background-color:white;"> <div style="left:0px; top:0px; width:600px; position:absolute"> <span typeof="mw:File"><a href="/wiki/File:Gene_structure_eukaryote_2_annotated.svg" title="File:Gene structure eukaryote 2 annotated.svg"><img alt="Eukaryote gene structure diagram" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Gene_structure_eukaryote_2_unannotated.svg/600px-Gene_structure_eukaryote_2_unannotated.svg.png" decoding="async" width="600" height="323" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Gene_structure_eukaryote_2_unannotated.svg/900px-Gene_structure_eukaryote_2_unannotated.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Gene_structure_eukaryote_2_unannotated.svg/1200px-Gene_structure_eukaryote_2_unannotated.svg.png 2x" data-file-width="1190" data-file-height="640" /></a></span> </div> <div style="text-align:left; font-size:11px; line-height:110%"> <div style="background-color:transparent; color:black"><div id="annotation_115x2" style="position:absolute; left:115px; top:2px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_473x2" style="position:absolute; left:473px; top:2px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_71x30" style="position:absolute; left:71px; top:30px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_71x41" style="position:absolute; left:71px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_164x41" style="position:absolute; left:164px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> </span></div> <div id="annotation_234x41" style="position:absolute; left:234px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/5%27UTR" class="mw-redirect" title="5'UTR">5'UTR </a> </span></div> <div id="annotation_320x41" style="position:absolute; left:320px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Open_reading_frame" title="Open reading frame">Open reading frame </a> </span></div> <div id="annotation_475x41" style="position:absolute; left:475px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/3%27UTR" class="mw-redirect" title="3'UTR">3'UTR </a> </span></div> <div id="annotation_535x30" style="position:absolute; left:535px; top:30px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_535x41" style="position:absolute; left:535px; top:41px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_145x67" style="position:absolute; left:145px; top:67px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Promoter_(genetics)#Elements" title="Promoter (genetics)">Proximal </a> </span></div> <div id="annotation_198x67" style="position:absolute; left:198px; top:67px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Promoter_(genetics)#Elements" title="Promoter (genetics)">Core </a> </span></div> <div id="annotation_268x67" style="position:absolute; left:268px; top:67px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Start_codon" title="Start codon">Start </a> </span></div> <div id="annotation_450x67" style="position:absolute; left:450px; top:67px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Stop_codon" title="Stop codon">Stop </a> </span></div> <div id="annotation_483x67" style="position:absolute; left:483px; top:67px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">Terminator</a> </span></div> <div id="annotation_51x120" style="position:absolute; left:51px; top:120px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription</a> </span></div> <div id="annotation_18x89" style="position:absolute; left:18px; top:89px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/DNA" title="DNA">DNA </a> </span></div> <div id="annotation_260x125" style="position:absolute; left:260px; top:125px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Exon" title="Exon">Exon </a> </span></div> <div id="annotation_341x125" style="position:absolute; left:341px; top:125px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Exon" title="Exon">Exon </a> </span></div> <div id="annotation_440x125" style="position:absolute; left:440px; top:125px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Exon" title="Exon">Exon </a> </span></div> <div id="annotation_314x138" style="position:absolute; left:314px; top:138px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Intron" title="Intron">Intron </a> </span></div> <div id="annotation_367x138" style="position:absolute; left:367px; top:138px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Intron" title="Intron">Intron </a> </span></div> <div id="annotation_51x171" style="position:absolute; left:51px; top:171px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcriptional<br />modification</a> </span></div> <div id="annotation_14x157" style="position:absolute; left:14px; top:157px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Primary_transcript" title="Primary transcript">Pre-<br />mRNA </a> </span></div> <div id="annotation_315x186" style="position:absolute; left:315px; top:186px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">Protein coding region </a> </span></div> <div id="annotation_250x198" style="position:absolute; left:250px; top:198px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/5%27cap" class="mw-redirect" title="5'cap">5'cap </a> </span></div> <div id="annotation_477x198" style="position:absolute; left:477px; top:198px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Poly-A_tail" class="mw-redirect" title="Poly-A tail">Poly-A tail </a> </span></div> <div id="annotation_51x248" style="position:absolute; left:51px; top:248px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">Translation</a> </span></div> <div id="annotation_14x221" style="position:absolute; left:14px; top:221px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">Mature<br />mRNA </a> </span></div> <div id="annotation_14x285" style="position:absolute; left:14px; top:285px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein" title="Protein">Protein </a> </span></div></div> </div> </div> <div class="thumbcaption" style="clear:left"><div style="float:left;margin-right:0.5em"><span typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/20px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/40px-Interactive_icon.svg.png 1.5x" data-file-width="133" data-file-height="200" /></span></span></div>The structure of a <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> protein-coding gene. <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence</a> controls when and where expression occurs for the <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">protein coding region</a> (red). <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions (yellow) regulate the <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of the gene into a pre-mRNA which is <a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">modified</a> to remove <a href="/wiki/Intron" title="Intron">introns</a> (light grey) and add a 5' cap and poly-A tail (dark grey). The mRNA <a href="/wiki/5%27_UTR" class="mw-redirect" title="5' UTR">5'</a> and <a href="/wiki/3%27_UTR" class="mw-redirect" title="3' UTR">3'</a> untranslated regions (blue) regulate <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> into the final protein product.<sup id="cite_ref-ShafeeLowe2017_57-0" class="reference"><a href="#cite_note-ShafeeLowe2017-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup></div> </div> </div> </td></tr></tbody></table> <table role="presentation" cellpadding="0" style="border-spacing:0; clear:right; float:right;"> <tbody><tr> <td><div class="thumb tright"> <div class="thumbinner" style="width:602px;"> <div class="skin-invert" style="position:relative; width:600px; height:300px; overflow:hidden; border:solid #ccc 1px; background-color:white;"> <div style="left:0px; top:0px; width:600px; position:absolute"> <span typeof="mw:File"><a href="/wiki/File:Gene_structure_prokaryote_2_annotated.svg" title="File:Gene structure prokaryote 2 annotated.svg"><img alt="Prokaryote gene structure diagram" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Gene_structure_prokaryote_2_unannotated.svg/960px-Gene_structure_prokaryote_2_unannotated.svg.png" decoding="async" width="600" height="295" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Gene_structure_prokaryote_2_unannotated.svg/1200px-Gene_structure_prokaryote_2_unannotated.svg.png 2x" data-file-width="1190" data-file-height="586" /></a></span> </div> <div style="text-align:left; font-size:11px; line-height:110%"> <div style="background-color:transparent; color:black"><div id="annotation_310x2" style="position:absolute; left:310px; top:2px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Operon" title="Operon">Polycistronic operon</a> </span></div> <div id="annotation_115x37" style="position:absolute; left:115px; top:37px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_475x37" style="position:absolute; left:475px; top:37px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence </a> </span></div> <div id="annotation_71x66" style="position:absolute; left:71px; top:66px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_535x66" style="position:absolute; left:535px; top:66px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">Enhancer</a> </span></div> <div id="annotation_535x77" style="position:absolute; left:535px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_71x77" style="position:absolute; left:71px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> /<a href="/wiki/Silencer_(genetics)" title="Silencer (genetics)">silencer</a> </span></div> <div id="annotation_139x77" style="position:absolute; left:139px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Operator_(genetics)" class="mw-redirect" title="Operator (genetics)">Operator</a> </span></div> <div id="annotation_187x77" style="position:absolute; left:187px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a> </span></div> <div id="annotation_236x77" style="position:absolute; left:236px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/5%27UTR" class="mw-redirect" title="5'UTR">5'UTR </a> </span></div> <div id="annotation_301x77" style="position:absolute; left:301px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Open_reading_frame" title="Open reading frame">ORF </a> </span></div> <div id="annotation_431x77" style="position:absolute; left:431px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Open_reading_frame" title="Open reading frame">ORF </a> </span></div> <div id="annotation_375x77" style="position:absolute; left:375px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Untranslated_region" title="Untranslated region">UTR </a> </span></div> <div id="annotation_475x77" style="position:absolute; left:475px; top:77px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/3%27UTR" class="mw-redirect" title="3'UTR">3'UTR </a> </span></div> <div id="annotation_268x103" style="position:absolute; left:268px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Start_codon" title="Start codon">Start </a> </span></div> <div id="annotation_410x103" style="position:absolute; left:410px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Start_codon" title="Start codon">Start </a> </span></div> <div id="annotation_344x103" style="position:absolute; left:344px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Stop_codon" title="Stop codon">Stop </a> </span></div> <div id="annotation_450x103" style="position:absolute; left:450px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Stop_codon" title="Stop codon">Stop </a> </span></div> <div id="annotation_483x103" style="position:absolute; left:483px; top:103px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">Terminator</a> </span></div> <div id="annotation_51x155" style="position:absolute; left:51px; top:155px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription</a> </span></div> <div id="annotation_18x124" style="position:absolute; left:18px; top:124px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/DNA" title="DNA">DNA </a> </span></div> <div id="annotation_239x170" style="position:absolute; left:239px; top:170px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Ribosome_binding_site" class="mw-redirect" title="Ribosome binding site">RBS </a> </span></div> <div id="annotation_382x170" style="position:absolute; left:382px; top:170px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Ribosome_binding_site" class="mw-redirect" title="Ribosome binding site">RBS </a> </span></div> <div id="annotation_264x156" style="position:absolute; left:264px; top:156px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">Protein coding region </a> </span></div> <div id="annotation_408x156" style="position:absolute; left:408px; top:156px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">Protein coding region </a> </span></div> <div id="annotation_14x195" style="position:absolute; left:14px; top:195px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA </a> </span></div> <div id="annotation_51x219" style="position:absolute; left:51px; top:219px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">Translation</a> </span></div> <div id="annotation_14x256" style="position:absolute; left:14px; top:256px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Protein" title="Protein">Protein </a> </span></div></div> </div> </div> <div class="thumbcaption" style="clear:left"><div style="float:left;margin-right:0.5em"><span typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/20px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/40px-Interactive_icon.svg.png 1.5x" data-file-width="133" data-file-height="200" /></span></span></div>The structure of a <a href="/wiki/Prokaryotic" class="mw-redirect" title="Prokaryotic">prokaryotic</a> <a href="/wiki/Operon" title="Operon">operon</a> of protein-coding genes. <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">Regulatory sequence</a> controls when expression occurs for the multiple <a href="/wiki/Protein_coding_region" class="mw-redirect" title="Protein coding region">protein coding regions</a> (red). <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a>, <a href="/wiki/Operator_(biology)" class="mw-redirect" title="Operator (biology)">operator</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a> regions (yellow) regulate the <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of the gene into an mRNA. The mRNA <a href="/wiki/Untranslated_region" title="Untranslated region">untranslated regions</a> (blue) regulate <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> into the final protein products.<sup id="cite_ref-ShafeeLowe2017_57-1" class="reference"><a href="#cite_note-ShafeeLowe2017-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup></div> </div> </div> </td></tr></tbody></table> </td></tr></tbody></table> <p>The <a href="/wiki/Gene_structure" title="Gene structure">structure of a protein-coding gene</a> consists of many elements of which the actual <a href="/wiki/Coding_region" title="Coding region">protein coding sequence</a> is often only a small part. These include introns and untranslated regions of the mature mRNA. Noncoding genes can also contain introns that are removed during processing to produce the mature functional RNA. </p><p>All genes are associated with <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">regulatory sequences</a> that are required for their expression. First, genes require a <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> sequence. The promoter is recognized and bound by <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> that recruit and help <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> bind to the region to initiate transcription.<sup id="cite_ref-MBOC_51-8" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7.1">: 7.1 </span></sup> The recognition typically occurs as a <a href="/wiki/Consensus_sequence" title="Consensus sequence">consensus sequence</a> like the <a href="/wiki/TATA_box" title="TATA box">TATA box</a>. A gene can have more than one promoter, resulting in messenger RNAs (<a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a>) that differ in how far they extend in the 5' end.<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> Highly transcribed genes have "strong" promoter sequences that form strong associations with transcription factors, thereby initiating transcription at a high rate. Others genes have "weak" promoters that form weak associations with transcription factors and initiate transcription less frequently.<sup id="cite_ref-MBOC_51-9" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7.2">: 7.2 </span></sup> <a href="/wiki/Eukaryote" title="Eukaryote">Eukaryotic</a> <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> regions are much more complex and difficult to identify than <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotic</a> promoters.<sup id="cite_ref-MBOC_51-10" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7.3">: 7.3 </span></sup> </p><p>Additionally, genes can have regulatory regions many kilobases upstream or downstream of the gene that alter expression. These act by <a href="/wiki/DNA_binding_site" title="DNA binding site">binding</a> to transcription factors which then cause the DNA to loop so that the regulatory sequence (and bound transcription factor) become close to the RNA polymerase binding site.<sup id="cite_ref-5questions_59-0" class="reference"><a href="#cite_note-5questions-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancers</a> increase transcription by binding an <a href="/wiki/Activator_(genetics)" title="Activator (genetics)">activator</a> protein which then helps to recruit the RNA polymerase to the promoter; conversely <a href="/wiki/Silencer_(DNA)" class="mw-redirect" title="Silencer (DNA)">silencers</a> bind <a href="/wiki/Repressor" title="Repressor">repressor</a> proteins and make the DNA less available for RNA polymerase.<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> </p><p>The mature messenger RNA produced from protein-coding genes contains <a href="/wiki/Untranslated_region" title="Untranslated region">untranslated regions</a> at both ends which contain binding sites for <a href="/wiki/Ribosome_binding_site" class="mw-redirect" title="Ribosome binding site">ribosomes</a>, <a href="/wiki/RNA-binding_protein" title="RNA-binding protein">RNA-binding proteins</a>, <a href="/wiki/MicroRNA" title="MicroRNA">miRNA</a>, as well as <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">terminator</a>, and <a href="/wiki/Start_codon" title="Start codon">start</a> and <a href="/wiki/Stop_codons" class="mw-redirect" title="Stop codons">stop codons</a>.<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> In addition, most eukaryotic <a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frames</a> contain untranslated <a href="/wiki/Intron" title="Intron">introns</a>, which are removed and <a href="/wiki/Exon" title="Exon">exons</a>, which are connected together in a process known as <a href="/wiki/RNA_splicing" title="RNA splicing">RNA splicing</a>. Finally, the ends of gene transcripts are defined by <a href="/wiki/Polyadenylation" title="Polyadenylation">cleavage and polyadenylation (CPA) sites</a>, where newly produced pre-mRNA gets cleaved and a string of ~200 adenosine monophosphates is added at the 3' end. The <a href="/wiki/Polyadenylation" title="Polyadenylation">poly(A)</a> tail protects mature mRNA from degradation and has other functions, affecting translation, localization, and transport of the transcript from the nucleus. Splicing, followed by CPA, generate the final <a href="/wiki/Mature_messenger_RNA" title="Mature messenger RNA">mature mRNA</a>, which encodes the protein or RNA product.<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> </p><p>Many noncoding genes in eukaryotes have different transcription termination mechanisms and they do not have poly(A) tails. </p><p>Many prokaryotic genes are organized into <a href="/wiki/Operon" title="Operon">operons</a>, with multiple protein-coding sequences that are transcribed as a unit.<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> The genes in an <a href="/wiki/Operon" title="Operon">operon</a> are transcribed as a continuous <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a>, referred to as a <a href="/wiki/Messenger_RNA#Monocistronic_versus_polycistronic_mRNA" title="Messenger RNA">polycistronic mRNA</a>. The term <a href="/wiki/Cistron" title="Cistron">cistron</a> in this context is equivalent to gene. The transcription of an operon's mRNA is often controlled by a <a href="/wiki/Repressor" title="Repressor">repressor</a> that can occur in an active or inactive state depending on the presence of specific metabolites.<sup id="cite_ref-pmid13718526_65-0" class="reference"><a href="#cite_note-pmid13718526-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> When active, the repressor binds to a DNA sequence at the beginning of the operon, called the <a href="/wiki/Operon#Operator" title="Operon">operator region</a>, and represses <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of the <a href="/wiki/Operon" title="Operon">operon</a>; when the repressor is inactive transcription of the operon can occur (see e.g. <a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a>). The products of operon genes typically have related functions and are involved in the same <a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">regulatory network</a>.<sup id="cite_ref-MBOC_51-11" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7.3">: 7.3 </span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Complexity">Complexity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=11" title="Edit section: Complexity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Though many genes have simple structures, as with much of biology, others can be quite complex or represent unusual edge-cases. Eukaryotic genes often have introns that are much larger than their exons,<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><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> and those introns can even have other genes <a href="/wiki/Nested_gene" title="Nested gene">nested inside them</a>.<sup id="cite_ref-Kumar_68-0" class="reference"><a href="#cite_note-Kumar-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> Associated enhancers may be many kilobase away, or even on entirely different chromosomes operating via physical contact between two chromosomes.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><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> A single gene can encode multiple different functional products by <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a>, and conversely a gene may be split across chromosomes but those transcripts are concatenated back together into a functional sequence by <a href="/wiki/Trans-splicing" title="Trans-splicing">trans-splicing</a>.<sup id="cite_ref-Lei2016_71-0" class="reference"><a href="#cite_note-Lei2016-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> It is also possible for <a href="/wiki/Overlapping_gene" title="Overlapping gene">overlapping genes</a> to share some of their DNA sequence, either on opposite strands or the same strand (in a different reading frame, or even the same reading frame).<sup id="cite_ref-Wright_2021_72-0" class="reference"><a href="#cite_note-Wright_2021-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Gene_expression">Gene expression</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=12" title="Edit section: Gene expression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div> <p>In all organisms, two steps are required to read the information encoded in a gene's DNA and produce the protein it specifies. First, the gene's DNA is <i><a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a></i> to messenger RNA (<a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a>).<sup id="cite_ref-MBOC_51-12" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6.1">: 6.1 </span></sup> Second, that mRNA is <i><a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translated</a></i> to protein.<sup id="cite_ref-MBOC_51-13" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6.2">: 6.2 </span></sup> RNA-coding genes must still go through the first step, but are not translated into protein.<sup id="cite_ref-Edd01_73-0" class="reference"><a href="#cite_note-Edd01-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> The process of producing a biologically functional molecule of either RNA or protein is called <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>, and the resulting molecule is called a <a href="/wiki/Gene_product" title="Gene product">gene product</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_code">Genetic code</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=13" title="Edit section: Genetic code"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:RNA-codons-aminoacids.svg" class="mw-file-description"><img alt="An RNA molecule consisting of nucleotides. Groups of three nucleotides are indicated as codons, with each corresponding to a specific amino acid." src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/RNA-codons-aminoacids.svg/330px-RNA-codons-aminoacids.svg.png" decoding="async" width="330" height="88" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/RNA-codons-aminoacids.svg/495px-RNA-codons-aminoacids.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6d/RNA-codons-aminoacids.svg/660px-RNA-codons-aminoacids.svg.png 2x" data-file-width="512" data-file-height="137" /></a><figcaption>Schematic of a single-stranded RNA molecule illustrating a series of three-base <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>. Each three-<a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> codon corresponds to an <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> when translated to protein.</figcaption></figure> <p>The nucleotide sequence of a gene's DNA specifies the amino acid sequence of a protein through the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>. Sets of three nucleotides, known as <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>, each correspond to a specific amino acid.<sup id="cite_ref-MBOC_51-14" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6">: 6 </span></sup> The principle that three sequential bases of DNA code for each amino acid was demonstrated in 1961 using frameshift mutations in the rIIB gene of bacteriophage T4<sup id="cite_ref-pmid13882203_74-0" class="reference"><a href="#cite_note-pmid13882203-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> (see <a href="/wiki/Crick,_Brenner_et_al._experiment" title="Crick, Brenner et al. experiment">Crick, Brenner et al. experiment</a>). </p><p>Additionally, a "<a href="/wiki/Start_codon" title="Start codon">start codon</a>", and three "<a href="/wiki/Stop_codon" title="Stop codon">stop codons</a>" indicate the beginning and end of the <a href="/wiki/Coding_region" title="Coding region">protein coding region</a>. There are 64 possible codons (four possible nucleotides at each of three positions, hence 4<sup>3</sup> possible codons) and only 20 standard amino acids; hence the code is redundant and multiple codons can specify the same amino acid. The correspondence between codons and amino acids is nearly universal among all known living organisms.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transcription">Transcription</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=14" title="Edit section: Transcription"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription</a> produces a single-stranded <a href="/wiki/RNA" title="RNA">RNA</a> molecule known as <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a>, whose nucleotide sequence is complementary to the DNA from which it was transcribed.<sup id="cite_ref-MBOC_51-15" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6.1">: 6.1 </span></sup> The mRNA acts as an intermediate between the DNA gene and its final protein product. The gene's DNA is used as a template to generate a <a href="/wiki/Base_pair" title="Base pair">complementary</a> mRNA. The mRNA matches the sequence of the gene's DNA <a href="/wiki/Coding_strand" title="Coding strand">coding strand</a> because it is synthesised as the complement of the <a href="/wiki/Template_strand" class="mw-redirect" title="Template strand">template strand</a>. Transcription is performed by an <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> called an <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>, which reads the template strand in the <a href="/wiki/3%27_end" class="mw-redirect" title="3' end">3'</a> to <a href="/wiki/5%27_end" class="mw-redirect" title="5' end">5'</a> direction and synthesizes the RNA from <a href="/wiki/5%27_end" class="mw-redirect" title="5' end">5'</a> to <a href="/wiki/3%27_end" class="mw-redirect" title="3' end">3'</a>. To initiate transcription, the polymerase first recognizes and binds a <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a> region of the gene. Thus, a major mechanism of <a href="/wiki/Gene_regulation" class="mw-redirect" title="Gene regulation">gene regulation</a> is the blocking or sequestering the promoter region, either by tight binding by <a href="/wiki/Repressor" title="Repressor">repressor</a> molecules that physically block the polymerase or by organizing the DNA so that the promoter region is not accessible.<sup id="cite_ref-MBOC_51-16" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7">: 7 </span></sup> </p><p>In <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>, transcription occurs in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>; for very long transcripts, translation may begin at the 5' end of the RNA while the 3' end is still being transcribed. In <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, transcription occurs in the nucleus, where the cell's DNA is stored. The RNA molecule produced by the polymerase is known as the <a href="/wiki/Primary_transcript" title="Primary transcript">primary transcript</a> and undergoes <a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">post-transcriptional modifications</a> before being exported to the cytoplasm for translation. One of the modifications performed is the <a href="/wiki/Splicing_(genetics)" class="mw-redirect" title="Splicing (genetics)">splicing</a> of <a href="/wiki/Intron" title="Intron">introns</a> which are sequences in the transcribed region that do not encode a protein. <a href="/wiki/Alternative_splicing" title="Alternative splicing">Alternative splicing</a> mechanisms can result in mature transcripts from the same gene having different sequences and thus coding for different proteins. This is a major form of regulation in eukaryotic cells and also occurs in some prokaryotes.<sup id="cite_ref-MBOC_51-17" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7.5">: 7.5 </span></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> <div class="mw-heading mw-heading3"><h3 id="Translation">Translation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=15" title="Edit section: Translation"><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:DNA_to_protein_or_ncRNA.svg" class="mw-file-description"><img alt="A protein-coding gene in DNA being transcribed and translated to a functional protein or a non-protein-coding gene being transcribed to a functional RNA" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/DNA_to_protein_or_ncRNA.svg/330px-DNA_to_protein_or_ncRNA.svg.png" decoding="async" width="330" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/DNA_to_protein_or_ncRNA.svg/500px-DNA_to_protein_or_ncRNA.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/DNA_to_protein_or_ncRNA.svg/960px-DNA_to_protein_or_ncRNA.svg.png 2x" data-file-width="1200" data-file-height="824" /></a><figcaption>Protein coding genes are transcribed to an <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> intermediate, then translated to a functional <a href="/wiki/Protein" title="Protein">protein</a>. RNA-coding genes are transcribed to a functional <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding RNA</a> (<span class="plainlinks"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a>: <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/3BSE">3BSE</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1OBB">1OBB</a>, <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/3TRA">3TRA</a></span>).</figcaption></figure> <p><a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">Translation</a> is the process by which a <a href="/wiki/Mature_messenger_RNA" title="Mature messenger RNA">mature mRNA</a> molecule is used as a template for synthesizing a new <a href="/wiki/Protein" title="Protein">protein</a>.<sup id="cite_ref-MBOC_51-18" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6.2">: 6.2 </span></sup> Translation is carried out by <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>, large complexes of RNA and protein responsible for carrying out the chemical reactions to add new <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> to a growing <a href="/wiki/Polypeptide_chain" class="mw-redirect" title="Polypeptide chain">polypeptide chain</a> by the formation of <a href="/wiki/Peptide_bond" title="Peptide bond">peptide bonds</a>. The genetic code is read three nucleotides at a time, in units called <a href="/wiki/Codon" class="mw-redirect" title="Codon">codons</a>, via interactions with specialized RNA molecules called <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNA</a> (tRNA). Each tRNA has three unpaired bases known as the <a href="/wiki/Anticodon" class="mw-redirect" title="Anticodon">anticodon</a> that are complementary to the codon it reads on the mRNA. The tRNA is also <a href="/wiki/Covalent" class="mw-redirect" title="Covalent">covalently</a> attached to the <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> specified by the complementary codon. When the tRNA binds to its complementary codon in an mRNA strand, the ribosome attaches its amino acid cargo to the new polypeptide chain, which is synthesized from <a href="/wiki/N-terminus" title="N-terminus">amino terminus</a> to <a href="/wiki/C-terminus" title="C-terminus">carboxyl terminus</a>. During and after synthesis, most new proteins must <a href="/wiki/Protein_folding" title="Protein folding">fold</a> to their active <a href="/wiki/Tertiary_structure" class="mw-redirect" title="Tertiary structure">three-dimensional structure</a> before they can carry out their cellular functions.<sup id="cite_ref-MBOC_51-19" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 3">: 3 </span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Regulation">Regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=16" title="Edit section: Regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Genes are regulated</a> so that they are <a href="/wiki/Gene_expression" title="Gene expression">expressed</a> only when the product is needed, since expression draws on limited resources.<sup id="cite_ref-MBOC_51-20" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 7">: 7 </span></sup> A cell regulates its gene expression depending on its <a href="/wiki/Environment_(biophysical)" class="mw-redirect" title="Environment (biophysical)">external environment</a> (e.g. <a href="/wiki/Nutrient" title="Nutrient">available nutrients</a>, <a href="/wiki/Heat_shock_protein" title="Heat shock protein">temperature</a> and other <a href="/wiki/Cellular_stress_response" title="Cellular stress response">stresses</a>), its internal environment (e.g. <a href="/wiki/Cell_division_cycle" class="mw-redirect" title="Cell division cycle">cell division cycle</a>, <a href="/wiki/Metabolism" title="Metabolism">metabolism</a>, <a href="/wiki/Infection" title="Infection">infection status</a>), and its <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">specific role</a> if in a <a href="/wiki/Multicellular" class="mw-redirect" title="Multicellular">multicellular</a> organism. Gene expression can be regulated at any step: from <a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">transcriptional initiation</a>, to <a href="/wiki/RNA_processing" class="mw-redirect" title="RNA processing">RNA processing</a>, to <a href="/wiki/Post-translational_modification" title="Post-translational modification">post-translational modification</a> of the protein. The regulation of <a href="/wiki/Lactose" title="Lactose">lactose</a> metabolism genes in <i><a href="/wiki/E._coli" class="mw-redirect" title="E. coli">E. coli</a></i> (<a href="/wiki/Lac_operon" title="Lac operon"><i>lac</i> operon</a>) was the first such mechanism to be described in 1961.<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> <div class="mw-heading mw-heading3"><h3 id="RNA_genes">RNA genes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=17" title="Edit section: RNA genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A typical protein-coding gene is first copied into <a href="/wiki/RNA" title="RNA">RNA</a> as an intermediate in the manufacture of the final protein product.<sup id="cite_ref-MBOC_51-21" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 6.1">: 6.1 </span></sup> In other cases, the RNA molecules are the actual functional products, as in the synthesis of <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a> and <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNA</a>. Some RNAs known as <a href="/wiki/Ribozyme" title="Ribozyme">ribozymes</a> are capable of <a href="/wiki/Enzyme" title="Enzyme">enzymatic function</a>, while others such as <a href="/wiki/MicroRNA" title="MicroRNA">microRNAs</a> and <a href="/wiki/Riboswitch" title="Riboswitch">riboswitches</a> have regulatory roles. The <a href="/wiki/DNA" title="DNA">DNA</a> sequences from which such RNAs are transcribed are known as <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding RNA genes</a>.<sup id="cite_ref-Edd01_73-1" class="reference"><a href="#cite_note-Edd01-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> </p><p>Some <a href="/wiki/Virus" title="Virus">viruses</a> store their entire genomes in the form of <a href="/wiki/RNA" title="RNA">RNA</a>, and contain no DNA at all.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> Because they use RNA to store genes, their <a href="/wiki/Cell_(biology)" title="Cell (biology)">cellular</a> <a href="/wiki/Host_(biology)" title="Host (biology)">hosts</a> may synthesize their proteins as soon as they are <a href="/wiki/Infection" title="Infection">infected</a> and without the delay in waiting for transcription.<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> On the other hand, RNA <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>, such as <a href="/wiki/HIV" title="HIV">HIV</a>, require the <a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">reverse transcription</a> of their <a href="/wiki/Genome" title="Genome">genome</a> from RNA into DNA before their proteins can be synthesized. </p> <div class="mw-heading mw-heading2"><h2 id="Inheritance">Inheritance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=18" title="Edit section: Inheritance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:Autosomal_recessive_-_mini.svg" class="mw-file-description"><img alt="Illustration of autosomal recessive inheritance. Each parent has one blue allele and one white allele. Each of their 4 children inherit one allele from each parent such that one child ends up with two blue alleles, one child has two white alleles and two children have one of each allele. Only the child with both blue alleles shows the trait because the trait is recessive." src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Autosomal_recessive_-_mini.svg/220px-Autosomal_recessive_-_mini.svg.png" decoding="async" width="220" height="286" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Autosomal_recessive_-_mini.svg/330px-Autosomal_recessive_-_mini.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/Autosomal_recessive_-_mini.svg/440px-Autosomal_recessive_-_mini.svg.png 2x" data-file-width="1201" data-file-height="1561" /></a><figcaption>Inheritance of a gene that has two different <a href="/wiki/Allele" title="Allele">alleles</a> (blue and white). The gene is located on an <a href="/wiki/Autosomal_chromosome" class="mw-redirect" title="Autosomal chromosome">autosomal chromosome</a>. The white allele is <a href="/wiki/Recessive" class="mw-redirect" title="Recessive">recessive</a> to the blue allele. The probability of each outcome in the children's generation is one quarter, or 25 percent.</figcaption></figure><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a> and <a href="/wiki/Heredity" title="Heredity">Heredity</a></div><p>Organisms inherit their genes from their parents. <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">Asexual</a> organisms simply inherit a complete copy of their parent's genome. <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">Sexual</a> organisms have two copies of each chromosome because they inherit one complete set from each parent.<sup id="cite_ref-MBOC_51-22" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 1">: 1 </span></sup> </p><div class="mw-heading mw-heading3"><h3 id="Mendelian_inheritance">Mendelian inheritance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=19" title="Edit section: Mendelian inheritance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>According to <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a>, variations in an organism's <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> (observable physical and behavioral characteristics) are due in part to variations in its <a href="/wiki/Genotype" title="Genotype">genotype</a> (particular set of genes). Each gene specifies a particular trait with a different sequence of a gene (<a href="/wiki/Allele" title="Allele">alleles</a>) giving rise to different phenotypes. Most eukaryotic organisms (such as the pea plants Mendel worked on) have two alleles for each trait, one inherited from each parent.<sup id="cite_ref-MBOC_51-23" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 20">: 20 </span></sup> </p><p>Alleles at a locus may be <a href="/wiki/Dominant_gene" class="mw-redirect" title="Dominant gene">dominant</a> or <a href="/wiki/Recessive_gene" class="mw-redirect" title="Recessive gene">recessive</a>; dominant alleles give rise to their corresponding phenotypes when paired with any other allele for the same trait, whereas recessive alleles give rise to their corresponding phenotype only when paired with another copy of the same allele. If you know the genotypes of the organisms, you can determine which alleles are dominant and which are recessive. For example, if the allele specifying tall stems in pea plants is dominant over the allele specifying short stems, then pea plants that inherit one tall allele from one parent and one short allele from the other parent will also have tall stems. Mendel's work demonstrated that alleles assort independently in the production of <a href="/wiki/Gamete" title="Gamete">gametes</a>, or <a href="/wiki/Germ_cell" title="Germ cell">germ cells</a>, ensuring variation in the next generation. Although Mendelian inheritance remains a good model for many traits determined by single genes (including a number of well-known <a href="/wiki/Genetic_disorder#Single-gene_disorder" title="Genetic disorder">genetic disorders</a>) it does not include the physical processes of DNA replication and cell division.<sup id="cite_ref-scitable_miko_81-0" class="reference"><a href="#cite_note-scitable_miko-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-scitable_chial_82-0" class="reference"><a href="#cite_note-scitable_chial-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="DNA_replication_and_cell_division">DNA replication and cell division</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=20" title="Edit section: DNA replication and cell division"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The growth, development, and reproduction of organisms relies on <a href="/wiki/Cell_division" title="Cell division">cell division</a>; the process by which a single <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> divides into two usually identical <a href="/wiki/Daughter_cell" class="mw-redirect" title="Daughter cell">daughter cells</a>. This requires first making a duplicate copy of every gene in the <a href="/wiki/Genome" title="Genome">genome</a> in a process called <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>.<sup id="cite_ref-MBOC_51-24" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 5.2">: 5.2 </span></sup> The copies are made by specialized <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> known as <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerases</a>, which "read" one strand of the double-helical DNA, known as the template strand, and synthesize a new complementary strand. Because the DNA double helix is held together by <a href="/wiki/Base_pair" title="Base pair">base pairing</a>, the sequence of one strand completely specifies the sequence of its complement; hence only one strand needs to be read by the enzyme to produce a faithful copy. The process of DNA replication is <a href="/wiki/Semiconservative_replication" title="Semiconservative replication">semiconservative</a>; that is, the copy of the genome inherited by each daughter cell contains one original and one newly synthesized strand of DNA.<sup id="cite_ref-MBOC_51-25" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 5.2">: 5.2 </span></sup> </p><p>The rate of DNA replication in living cells was first measured as the rate of phage T4 DNA elongation in phage-infected <i>E. coli</i> and found to be impressively rapid.<sup id="cite_ref-pmid789903_83-0" class="reference"><a href="#cite_note-pmid789903-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> During the period of exponential DNA increase at 37 °C, the rate of elongation was 749 nucleotides per second. </p><p>After DNA replication, the cell must physically separate the two genome copies and divide into two distinct membrane-bound cells.<sup id="cite_ref-MBOC_51-26" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 18.2">: 18.2 </span></sup> In <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> (<a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>) this usually occurs via a relatively simple process called <a href="/wiki/Binary_fission" class="mw-redirect" title="Binary fission">binary fission</a>, in which each circular genome attaches to the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> and is separated into the daughter cells as the membrane <a href="/wiki/Invagination" title="Invagination">invaginates</a> to split the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> into two membrane-bound portions. Binary fission is extremely fast compared to the rates of cell division in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>. Eukaryotic cell division is a more complex process known as the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>; DNA replication occurs during a phase of this cycle known as <a href="/wiki/S_phase" title="S phase">S phase</a>, whereas the process of segregating <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> and splitting the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> occurs during <a href="/wiki/M_phase" class="mw-redirect" title="M phase">M phase</a>.<sup id="cite_ref-MBOC_51-27" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 18.1">: 18.1 </span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Molecular_inheritance">Molecular inheritance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=21" title="Edit section: Molecular inheritance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The duplication and transmission of genetic material from one generation of cells to the next is the basis for molecular inheritance and the link between the classical and molecular pictures of genes. Organisms inherit the characteristics of their parents because the cells of the offspring contain copies of the genes in their parents' cells. In <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexually reproducing</a> organisms, the offspring will be a genetic copy or <a href="/wiki/Clone_(genetics)" class="mw-redirect" title="Clone (genetics)">clone</a> of the parent organism. In <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexually reproducing</a> organisms, a specialized form of cell division called <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> produces cells called <a href="/wiki/Gamete" title="Gamete">gametes</a> or <a href="/wiki/Germ_cell" title="Germ cell">germ cells</a> that are <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a>, or contain only one copy of each gene.<sup id="cite_ref-MBOC_51-28" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 20.2">: 20.2 </span></sup> The gametes produced by females are called <a href="/wiki/Egg_(biology)" class="mw-redirect" title="Egg (biology)">eggs</a> or ova, and those produced by males are called <a href="/wiki/Sperm" title="Sperm">sperm</a>. Two gametes fuse to form a <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a> <a href="/wiki/Fertilized_egg" class="mw-redirect" title="Fertilized egg">fertilized egg</a>, a single cell that has two sets of genes, with one copy of each gene from the mother and one from the father.<sup id="cite_ref-MBOC_51-29" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 20">: 20 </span></sup> </p><p>During the process of meiotic cell division, an event called <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> or <i>crossing-over</i> can sometimes occur, in which a length of DNA on one <a href="/wiki/Chromatid" title="Chromatid">chromatid</a> is swapped with a length of DNA on the corresponding homologous non-sister chromatid. This can result in reassortment of otherwise linked alleles.<sup id="cite_ref-MBOC_51-30" class="reference"><a href="#cite_note-MBOC-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 5.5">: 5.5 </span></sup> The Mendelian principle of independent assortment asserts that each of a parent's two genes for each trait will sort independently into gametes; which allele an organism inherits for one trait is unrelated to which allele it inherits for another trait. This is in fact only true for genes that do not reside on the same chromosome or are located very far from one another on the same chromosome. The closer two genes lie on the same chromosome, the more closely they will be associated in gametes and the more often they will appear together (known as <a href="/wiki/Genetic_linkage" title="Genetic linkage">genetic linkage</a>).<sup id="cite_ref-scitable_lobo_84-0" class="reference"><a href="#cite_note-scitable_lobo-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> Genes that are very close are essentially never separated because it is extremely unlikely that a crossover point will occur between them.<sup id="cite_ref-scitable_lobo_84-1" class="reference"><a href="#cite_note-scitable_lobo-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genome">Genome</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=22" title="Edit section: Genome"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Genome" title="Genome">genome</a> is the total genetic material of an organism and includes both the genes and <a href="/wiki/Non-coding_sequence" class="mw-redirect" title="Non-coding sequence">non-coding sequences</a>.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> Eukaryotic genes can be annotated using FINDER.<sup id="cite_ref-pmid33879057_86-0" class="reference"><a href="#cite_note-pmid33879057-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Number_of_genes">Number of genes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=23" title="Edit section: Number of genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:Gene_numbers.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/89/Gene_numbers.svg/960px-Gene_numbers.svg.png" decoding="async" width="560" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/89/Gene_numbers.svg/1120px-Gene_numbers.svg.png 2x" data-file-width="1200" data-file-height="376" /></a><figcaption>Depiction of numbers of genes for representative <a href="/wiki/Plant" title="Plant">plants</a> (green), <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> (blue), <a href="/wiki/Invertebrate" title="Invertebrate">invertebrates</a> (orange), <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> (yellow), <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> (purple), and <a href="/wiki/Virus" title="Virus">viruses</a> (grey). An inset on the right shows the smaller genomes expanded 100-fold area-wise.<sup id="cite_ref-WatsonJD_87-0" class="reference"><a href="#cite_note-WatsonJD-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11935017_91-0" class="reference"><a href="#cite_note-pmid11935017-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-mitochondrial_genome_92-0" class="reference"><a href="#cite_note-mitochondrial_genome-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ReferenceB_94-0" class="reference"><a href="#cite_note-ReferenceB-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The <a href="/wiki/Genome_size" title="Genome size">genome size</a>, and the number of genes it encodes varies widely between organisms. The smallest genomes occur in <a href="/wiki/Virus" title="Virus">viruses</a>,<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Viroid" title="Viroid">viroids</a> (which act as a single non-coding RNA gene).<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> Conversely, plants can have extremely large genomes,<sup id="cite_ref-97" class="reference"><a href="#cite_note-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> with <a href="/wiki/Oryza_sativa" title="Oryza sativa">rice</a> containing >46,000 protein-coding genes.<sup id="cite_ref-pmid11935017_91-1" class="reference"><a href="#cite_note-pmid11935017-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> The total number of protein-coding genes (the Earth's <a href="/wiki/Proteome" title="Proteome">proteome</a>) is estimated to be 5 million sequences.<sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> </p><p>Although the number of base-pairs of DNA in the human genome has been known since the 1950s, the estimated number of genes has changed over time as definitions of genes, and methods of detecting them have been refined. Initial theoretical predictions of the number of human genes in the 1960s and 1970s were based on mutation load estimates and the numbers of mRNAs and these estimates tended to be about 30,000 protein-coding genes.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> During the 1990s there were guesstimates of up to 100,000 genes and early data on detection of mRNAs (<a href="/wiki/Expressed_sequence_tag" title="Expressed sequence tag">expressed sequence tags</a>) suggested more than the traditional value of 30,000 genes that had been reported in the textbooks during the 1980s.<sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p><p>The initial draft sequences of the human genome confirmed the earlier predictions of about 30,000 protein-coding genes however that estimate has fallen to about 19,000 with the ongoing <a href="/wiki/GENCODE" title="GENCODE">GENCODE</a> annotation project.<sup id="cite_ref-103" class="reference"><a href="#cite_note-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> The number of noncoding genes is not known with certainty but the latest estimates from Ensembl suggest 26,000 noncoding genes.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Essential_genes">Essential genes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=24" title="Edit section: Essential genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Essential_gene" title="Essential gene">Essential gene</a></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:Syn3_genome.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Syn3_genome.svg/300px-Syn3_genome.svg.png" decoding="async" width="300" height="179" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Syn3_genome.svg/450px-Syn3_genome.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Syn3_genome.svg/600px-Syn3_genome.svg.png 2x" data-file-width="340" data-file-height="203" /></a><figcaption>Gene functions in the minimal <a href="/wiki/Genome" title="Genome">genome</a> of the <a href="/wiki/Synthetic_biology" title="Synthetic biology">synthetic organism</a>, <i><a href="/wiki/Syn_3" class="mw-redirect" title="Syn 3">Syn 3</a></i><sup id="cite_ref-Hutchison_105-0" class="reference"><a href="#cite_note-Hutchison-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Essential genes are the set of genes thought to be critical for an organism's survival.<sup id="cite_ref-106" class="reference"><a href="#cite_note-106"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> This definition assumes the abundant availability of all relevant <a href="/wiki/Nutrient" title="Nutrient">nutrients</a> and the absence of environmental stress. Only a small portion of an organism's genes are essential. In bacteria, an estimated 250–400 genes are essential for <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i> and <i><a href="/wiki/Bacillus_subtilis" title="Bacillus subtilis">Bacillus subtilis</a></i>, which is less than 10% of their genes.<sup id="cite_ref-107" class="reference"><a href="#cite_note-107"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-108" class="reference"><a href="#cite_note-108"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_109-0" class="reference"><a href="#cite_note-:0-109"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> Half of these genes are <a href="/wiki/Ortholog" class="mw-redirect" title="Ortholog">orthologs</a> in both organisms and are largely involved in <a href="/wiki/Protein_synthesis" class="mw-redirect" title="Protein synthesis">protein synthesis</a>.<sup id="cite_ref-:0_109-1" class="reference"><a href="#cite_note-:0-109"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> In the budding yeast <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> the number of essential genes is slightly higher, at 1000 genes (~20% of their genes).<sup id="cite_ref-ReferenceA_110-0" class="reference"><a href="#cite_note-ReferenceA-110"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> Although the number is more difficult to measure in higher eukaryotes, mice and humans are estimated to have around 2000 essential genes (~10% of their genes).<sup id="cite_ref-georgi_111-0" class="reference"><a href="#cite_note-georgi-111"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> The synthetic organism, <i><a href="/wiki/Syn_3" class="mw-redirect" title="Syn 3">Syn 3</a></i>, has a minimal genome of 473 essential genes and quasi-essential genes (necessary for fast growth), although 149 have unknown function.<sup id="cite_ref-Hutchison_105-1" class="reference"><a href="#cite_note-Hutchison-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> </p><p>Essential genes include <a href="/wiki/Housekeeping_gene" title="Housekeeping gene">housekeeping genes</a> (critical for basic cell functions)<sup id="cite_ref-112" class="reference"><a href="#cite_note-112"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> as well as genes that are expressed at different times in the organisms <a href="/wiki/Developmental_biology" title="Developmental biology">development</a> or <a href="/wiki/Biological_life_cycle" title="Biological life cycle">life cycle</a>.<sup id="cite_ref-113" class="reference"><a href="#cite_note-113"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup> Housekeeping genes are used as <a href="/wiki/Experimental_control" class="mw-redirect" title="Experimental control">experimental controls</a> when <a href="/wiki/Gene_expression_analysis" class="mw-redirect" title="Gene expression analysis">analysing gene expression</a>, since they are <a href="/wiki/Gene_expression" title="Gene expression">constitutively expressed</a> at a relatively constant level. </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_and_genomic_nomenclature">Genetic and genomic nomenclature</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=25" title="Edit section: Genetic and genomic nomenclature"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Gene_nomenclature" title="Gene nomenclature">Gene nomenclature</a> was established by the <a href="/wiki/HUGO_Gene_Nomenclature_Committee" title="HUGO Gene Nomenclature Committee">HUGO Gene Nomenclature Committee</a> (HGNC), a committee of the <a href="/wiki/Human_Genome_Organisation" title="Human Genome Organisation">Human Genome Organisation</a>, for each known human gene in the form of an approved gene name and <a href="/wiki/Symbol" title="Symbol">symbol</a> (short-form <a href="/wiki/Abbreviation" title="Abbreviation">abbreviation</a>), which can be accessed through a database maintained by HGNC. Symbols are chosen to be unique, and each gene has only one symbol (although approved symbols sometimes change). Symbols are preferably kept consistent with other members of a <a href="/wiki/Gene_family" title="Gene family">gene family</a> and with homologs in other species, particularly the <a href="/wiki/Mouse" title="Mouse">mouse</a> due to its role as a common <a href="/wiki/Model_organism" title="Model organism">model organism</a>.<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetic_engineering">Genetic engineering</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=26" title="Edit section: Genetic engineering"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:Breeding_transgenesis_cisgenesis.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Breeding_transgenesis_cisgenesis.svg/330px-Breeding_transgenesis_cisgenesis.svg.png" decoding="async" width="300" height="257" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Breeding_transgenesis_cisgenesis.svg/500px-Breeding_transgenesis_cisgenesis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/80/Breeding_transgenesis_cisgenesis.svg/960px-Breeding_transgenesis_cisgenesis.svg.png 2x" data-file-width="1077" data-file-height="921" /></a><figcaption>Comparison of conventional plant breeding with transgenic and cisgenic genetic modification</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></div> <p>Genetic engineering is the modification of an organism's <a href="/wiki/Genome" title="Genome">genome</a> through <a href="/wiki/Biotechnology" title="Biotechnology">biotechnology</a>. Since the 1970s, a <a href="/wiki/Genetic_engineering_techniques" title="Genetic engineering techniques">variety of techniques</a> have been developed to specifically add, remove and edit genes in an organism.<sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup> Recently developed <a href="/wiki/Genome_engineering" class="mw-redirect" title="Genome engineering">genome engineering</a> techniques use engineered <a href="/wiki/Nuclease" title="Nuclease">nuclease</a> <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> to create targeted <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> in a <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> to either disrupt or edit a gene when the break is repaired.<sup id="cite_ref-116" class="reference"><a href="#cite_note-116"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ran_119-0" class="reference"><a href="#cite_note-ran-119"><span class="cite-bracket">[</span>119<span class="cite-bracket">]</span></a></sup> The related term <a href="/wiki/Synthetic_biology" title="Synthetic biology">synthetic biology</a> is sometimes used to refer to extensive genetic engineering of an organism.<sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">[</span>120<span class="cite-bracket">]</span></a></sup> </p><p>Genetic engineering is now a routine research tool with <a href="/wiki/Model_organism" title="Model organism">model organisms</a>. For example, genes are easily added to <a href="/wiki/Bacteria" title="Bacteria">bacteria</a><sup id="cite_ref-121" class="reference"><a href="#cite_note-121"><span class="cite-bracket">[</span>121<span class="cite-bracket">]</span></a></sup> and lineages of <a href="/wiki/Knockout_mice" class="mw-redirect" title="Knockout mice">knockout mice</a> with a specific gene's function disrupted are used to investigate that gene's function.<sup id="cite_ref-122" class="reference"><a href="#cite_note-122"><span class="cite-bracket">[</span>122<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-123" class="reference"><a href="#cite_note-123"><span class="cite-bracket">[</span>123<span class="cite-bracket">]</span></a></sup> Many organisms have been genetically modified for applications in <a href="/wiki/Agriculture" title="Agriculture">agriculture</a>, industrial biotechnology, and <a href="/wiki/Medicine" title="Medicine">medicine</a>. </p><p>For multicellular organisms, typically the <a href="/wiki/Embryo" title="Embryo">embryo</a> is engineered which grows into the adult <a href="/wiki/Genetically_modified_organism" title="Genetically modified organism">genetically modified organism</a>.<sup id="cite_ref-Deng_2007_124-0" class="reference"><a href="#cite_note-Deng_2007-124"><span class="cite-bracket">[</span>124<span class="cite-bracket">]</span></a></sup> However, the genomes of cells in an adult organism can be edited using <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a> techniques to treat genetic diseases. </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=Gene&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:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 22em;"> <ul><li><a href="/wiki/Biological_patent" title="Biological patent">Biological patent</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centric view of evolution</a>      </li> <li><a href="/wiki/Gene_dosage" title="Gene dosage">Gene dosage</a></li> <li><a href="/wiki/Gene_redundancy" title="Gene redundancy">Gene redundancy</a></li> <li><a href="/wiki/Gene_silencing" title="Gene silencing">Gene silencing</a></li> <li><a href="/wiki/Genetic_algorithm" title="Genetic algorithm">Genetic algorithm</a></li> <li><a href="/wiki/Haplotype" title="Haplotype">Haplotype</a></li> <li><a href="/wiki/List_of_gene_prediction_software" title="List of gene prediction software">List of gene prediction software</a></li> <li><a href="/wiki/Lists_of_human_genes" title="Lists of human genes">Lists of human genes</a></li> <li><a href="/wiki/Predictive_medicine" title="Predictive medicine">Predictive medicine</a></li> <li><a href="/wiki/Quantitative_trait_locus" title="Quantitative trait locus">Quantitative trait locus</a></li> <li><a href="/wiki/Selfish_genetic_element" title="Selfish genetic element">Selfish genetic element</a></li> <li><a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">Whole genome sequencing</a></li></ul></div> <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=Gene&action=edit&section=28" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Citations">Citations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=29" title="Edit section: Citations"><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-mendgene2-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-mendgene2_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-mendgene2_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-mendgene2_1-2"><sup><i><b>c</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="CITEREFOrgogozoPeluffoMorizot2016" class="citation journal cs1">Orgogozo V, Peluffo AE, Morizot B (2016). <a rel="nofollow" class="external text" href="https://hal.archives-ouvertes.fr/hal-01354346/file/Orgogozo2016-gene.pdf">"The 'Mendelian Gene' and the 'Molecular Gene': Two Relevant Concepts of Genetic Units"</a> <span class="cs1-format">(PDF)</span>. <i>Current Topics in Developmental Biology</i>. <b>119</b>: <span class="nowrap">1–</span>26. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fbs.ctdb.2016.03.002">10.1016/bs.ctdb.2016.03.002</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/27282022">27282022</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Topics+in+Developmental+Biology&rft.atitle=The+%27Mendelian+Gene%27+and+the+%27Molecular+Gene%27%3A+Two+Relevant+Concepts+of+Genetic+Units&rft.volume=119&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1-%3C%2Fspan%3E26&rft.date=2016&rft_id=info%3Adoi%2F10.1016%2Fbs.ctdb.2016.03.002&rft_id=info%3Apmid%2F27282022&rft.aulast=Orgogozo&rft.aufirst=V&rft.au=Peluffo%2C+AE&rft.au=Morizot%2C+B&rft_id=https%3A%2F%2Fhal.archives-ouvertes.fr%2Fhal-01354346%2Ffile%2FOrgogozo2016-gene.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-MedlinePlus_2020-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-MedlinePlus_2020_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://medlineplus.gov/genetics/understanding/basics/gene/">"What is a gene?: MedlinePlus Genetics"</a>. <i>MedlinePlus</i>. 17 September 2020. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200516195548/http://ghr.nlm.nih.gov/handbook/basics/gene">Archived</a> from the original on 16 May 2020<span class="reference-accessdate">. Retrieved <span class="nowrap">4 January</span> 2021</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=MedlinePlus&rft.atitle=What+is+a+gene%3F%3A+MedlinePlus+Genetics&rft.date=2020-09-17&rft_id=https%3A%2F%2Fmedlineplus.gov%2Fgenetics%2Funderstanding%2Fbasics%2Fgene%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-Hirsch_2002_p.-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Hirsch_2002_p._3-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFHirsch2002" class="citation book cs1">Hirsch ED (2002). <i>The new dictionary of cultural literacy</i>. 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Series A, Mathematical, Physical, and Engineering Sciences</i>. <b>366</b> (1878): <span class="nowrap">3001–</span>15. <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/2008RSPTA.366.3001N">2008RSPTA.366.3001N</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/10.1098%2Frsta.2008.0086">10.1098/rsta.2008.0086</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/18559318">18559318</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions.+Series+A%2C+Mathematical%2C+Physical%2C+and+Engineering+Sciences&rft.atitle=Genes+and+causation&rft.volume=366&rft.issue=1878&rft.pages=%3Cspan+class%3D%22nowrap%22%3E3001-%3C%2Fspan%3E15&rft.date=2008-09&rft_id=info%3Apmid%2F18559318&rft_id=info%3Adoi%2F10.1098%2Frsta.2008.0086&rft_id=info%3Abibcode%2F2008RSPTA.366.3001N&rft.aulast=Noble&rft.aufirst=D&rft_id=https%3A%2F%2Fdoi.org%2F10.1098%252Frsta.2008.0086&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-31"><span class="mw-cite-backlink"><b><a href="#cite_ref-31">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFMagnello2013" class="citation book cs1">Magnello, M.E. 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Retrieved <span class="nowrap">6 May</span> 2007</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Intracellulare+Pangenese&rft.place=Jena&rft.pub=Verlag+von+Gustav+Fischer&rft.date=1889&rft.aulast=de+Vries&rft.aufirst=H&rft_id=http%3A%2F%2Fwww.esp.org%2Fbooks%2Fdevries%2Fpangenesis%2Ffacsimile%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span> Translated in 1908 from German to English by Open Court Publishing Co., Chicago, 1910</span> </li> <li id="cite_note-Johannsen-36"><span class="mw-cite-backlink"><b><a href="#cite_ref-Johannsen_36-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFJohannsen1909" class="citation book cs1 cs1-prop-foreign-lang-source">Johannsen W (1909). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/15717#page/134/mode/1up"><i>Elemente der exakten Erblichkeitslehre</i></a> [<i>Elements of the exact theory of heredity</i>] (in German). Jena, Germany: Gustav Fischer. p. 124. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210624195335/https://www.biodiversitylibrary.org/item/15717#page/134/mode/1up">Archived</a> from the original on 24 June 2021<span class="reference-accessdate">. Retrieved <span class="nowrap">16 June</span> 2021</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Elemente+der+exakten+Erblichkeitslehre&rft.place=Jena%2C+Germany&rft.pages=124&rft.pub=Gustav+Fischer&rft.date=1909&rft.aulast=Johannsen&rft.aufirst=W&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F15717%23page%2F134%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span> From p. 124: <i>"Dieses "etwas" in den Gameten bezw. in der Zygote, ... – kurz, was wir eben Gene nennen wollen – bedingt sind."</i> (This "something" in the gametes or in the zygote, which has crucial importance for the character of the organism, is usually called by the quite ambiguous term <i>Anlagen</i> [primordium, from the German word <i>Anlage</i> for "plan, arrangement; rough sketch"]. Many other terms have been suggested, mostly unfortunately in closer connection with certain hypothetical opinions. The word "pangene", which was introduced by Darwin, is perhaps used most frequently in place of <i>Anlagen</i>. However, the word "pangene" was not well chosen, as it is a compound word containing the roots <i>pan</i> (the neuter form of Πας all, every) and <i>gen</i> (from γί-γ(ε)ν-ομαι, to become). Only the meaning of this latter [i.e., <i>gen</i>] comes into consideration here; just the basic idea – [namely,] that a trait in the developing organism can be determined or is influenced by "something" in the gametes – should find expression. No hypothesis about the nature of this "something" should be postulated or supported by it. For that reason it seems simplest to use in isolation the last syllable <i>gen</i> from Darwin's well-known word, which alone is of interest to us, in order to replace, with it, the poor, ambiguous word <i>Anlage</i>. Thus we will say simply "gene" and "genes" for "pangene" and "pangenes". The word gene is completely free of any hypothesis; it expresses only the established fact that in any case many traits of the organism are determined by specific, separable, and thus independent "conditions", "foundations", "plans" – in short, precisely what we want to call genes.)</span> </li> <li id="cite_note-37"><span class="mw-cite-backlink"><b><a href="#cite_ref-37">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFBateson1906" class="citation book cs1">Bateson W (1906). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/206746#page/129/mode/1up">"The progress of genetic research"</a>. In Wilks W (ed.). <i>Report of the Third International Conference 1906 on Genetics</i>. London, England: Royal Horticultural Society. pp. <span class="nowrap">90–</span>97. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20230627221528/https://www.biodiversitylibrary.org/item/206746#page/129/mode/1up">Archived</a> from the original on 27 June 2023<span class="reference-accessdate">. Retrieved <span class="nowrap">17 June</span> 2021</span>. <q>... the science itself [i.e. the study of the breeding and hybridisation of plants] is still nameless, and we can only describe our pursuit by cumbrous and often misleading periphrasis. To meet this difficulty I suggest for the consideration of this Congress the term <i>Genetics</i>, which sufficiently indicates that our labors are devoted to the elucidation of the phenomena of heredity and variation: in other words, to the physiology of Descent, with implied bearing on the theoretical problems of the evolutionist and the systematist, and application to the practical problems of breeders, whether of animals or plants.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=The+progress+of+genetic+research&rft.btitle=Report+of+the+Third+International+Conference+1906+on+Genetics&rft.place=London%2C+England&rft.pages=%3Cspan+class%3D%22nowrap%22%3E90-%3C%2Fspan%3E97&rft.pub=Royal+Horticultural+Society&rft.date=1906&rft.aulast=Bateson&rft.aufirst=W&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F206746%23page%2F129%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-Avery_et_al-38"><span class="mw-cite-backlink"><b><a href="#cite_ref-Avery_et_al_38-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAveryMacleodMcCarty1944" class="citation journal cs1">Avery OT, Macleod CM, McCarty M (February 1944). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2135445">"Studies on the Chemical Nature of the Substance Inducing Transformation of Pneumococcal Types: Induction of Transformation by a Desoxyribonucleic Acid Fraction Isolated From Pneumococcus Type III"</a>. <i>The Journal of Experimental Medicine</i>. <b>79</b> (2): <span class="nowrap">137–</span>58. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1084%2Fjem.79.2.137">10.1084/jem.79.2.137</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/PMC2135445">2135445</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/19871359">19871359</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+Experimental+Medicine&rft.atitle=Studies+on+the+Chemical+Nature+of+the+Substance+Inducing+Transformation+of+Pneumococcal+Types%3A+Induction+of+Transformation+by+a+Desoxyribonucleic+Acid+Fraction+Isolated+From+Pneumococcus+Type+III&rft.volume=79&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E137-%3C%2Fspan%3E58&rft.date=1944-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2135445%23id-name%3DPMC&rft_id=info%3Apmid%2F19871359&rft_id=info%3Adoi%2F10.1084%2Fjem.79.2.137&rft.aulast=Avery&rft.aufirst=OT&rft.au=Macleod%2C+CM&rft.au=McCarty%2C+M&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2135445&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span> Reprint: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAveryMacLeodMcCarty1979" class="citation journal cs1">Avery OT, MacLeod CM, McCarty M (February 1979). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2184805">"Studies on the chemical nature of the substance inducing transformation of pneumococcal types. Inductions of transformation by a desoxyribonucleic acid fraction isolated from pneumococcus type III"</a>. <i>The Journal of Experimental Medicine</i>. <b>149</b> (2): <span class="nowrap">297–</span>326. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1084%2Fjem.149.2.297">10.1084/jem.149.2.297</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/PMC2184805">2184805</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/33226">33226</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+Experimental+Medicine&rft.atitle=Studies+on+the+chemical+nature+of+the+substance+inducing+transformation+of+pneumococcal+types.+Inductions+of+transformation+by+a+desoxyribonucleic+acid+fraction+isolated+from+pneumococcus+type+III&rft.volume=149&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E297-%3C%2Fspan%3E326&rft.date=1979-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2184805%23id-name%3DPMC&rft_id=info%3Apmid%2F33226&rft_id=info%3Adoi%2F10.1084%2Fjem.149.2.297&rft.aulast=Avery&rft.aufirst=OT&rft.au=MacLeod%2C+CM&rft.au=McCarty%2C+M&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2184805&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-39"><span class="mw-cite-backlink"><b><a href="#cite_ref-39">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFHersheyChase1952" class="citation journal cs1">Hershey AD, Chase M (May 1952). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2147348">"Independent functions of viral protein and nucleic acid in growth of bacteriophage"</a>. <i>The Journal of General Physiology</i>. <b>36</b> (1): <span class="nowrap">39–</span>56. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1085%2Fjgp.36.1.39">10.1085/jgp.36.1.39</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/PMC2147348">2147348</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/12981234">12981234</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+Physiology&rft.atitle=Independent+functions+of+viral+protein+and+nucleic+acid+in+growth+of+bacteriophage&rft.volume=36&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E39-%3C%2Fspan%3E56&rft.date=1952-05&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2147348%23id-name%3DPMC&rft_id=info%3Apmid%2F12981234&rft_id=info%3Adoi%2F10.1085%2Fjgp.36.1.39&rft.aulast=Hershey&rft.aufirst=AD&rft.au=Chase%2C+M&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2147348&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-40"><span class="mw-cite-backlink"><b><a href="#cite_ref-40">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFJudson1979" class="citation book cs1"><a href="/wiki/Horace_Freeland_Judson" title="Horace Freeland Judson">Judson H</a> (1979). <i>The Eighth Day of Creation: Makers of the Revolution in Biology</i>. Cold Spring Harbor Laboratory Press. pp. <span class="nowrap">51–</span>169. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-477-7" title="Special:BookSources/978-0-87969-477-7"><bdi>978-0-87969-477-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Eighth+Day+of+Creation%3A+Makers+of+the+Revolution+in+Biology&rft.pages=%3Cspan+class%3D%22nowrap%22%3E51-%3C%2Fspan%3E169&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=1979&rft.isbn=978-0-87969-477-7&rft.aulast=Judson&rft.aufirst=H&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-watsoncrick_1953a-41"><span class="mw-cite-backlink"><b><a href="#cite_ref-watsoncrick_1953a_41-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWatsonCrick1953" class="citation journal cs1">Watson JD, Crick FH (April 1953). "Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid". <i>Nature</i>. <b>171</b> (4356): <span class="nowrap">737–</span>8. <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/1953Natur.171..737W">1953Natur.171..737W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F171737a0">10.1038/171737a0</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/13054692">13054692</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Molecular+Structure+of+Nucleic+Acids%3A+A+Structure+for+Deoxyribose+Nucleic+Acid&rft.volume=171&rft.issue=4356&rft.pages=%3Cspan+class%3D%22nowrap%22%3E737-%3C%2Fspan%3E8&rft.date=1953-04&rft_id=info%3Apmid%2F13054692&rft_id=info%3Adoi%2F10.1038%2F171737a0&rft_id=info%3Abibcode%2F1953Natur.171..737W&rft.aulast=Watson&rft.aufirst=JD&rft.au=Crick%2C+FH&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-pmid16589677-42"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid16589677_42-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFBenzer1955" class="citation journal cs1">Benzer S (June 1955). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC528093">"Fine Structure of a Genetic Region in Bacteriophage"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>41</b> (6): <span class="nowrap">344–</span>54. <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/1955PNAS...41..344B">1955PNAS...41..344B</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.41.6.344">10.1073/pnas.41.6.344</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/PMC528093">528093</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/16589677">16589677</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=Fine+Structure+of+a+Genetic+Region+in+Bacteriophage&rft.volume=41&rft.issue=6&rft.pages=%3Cspan+class%3D%22nowrap%22%3E344-%3C%2Fspan%3E54&rft.date=1955-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC528093%23id-name%3DPMC&rft_id=info%3Apmid%2F16589677&rft_id=info%3Adoi%2F10.1073%2Fpnas.41.6.344&rft_id=info%3Abibcode%2F1955PNAS...41..344B&rft.aulast=Benzer&rft.aufirst=S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC528093&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-pmid16590553-43"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid16590553_43-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFBenzer1959" class="citation journal cs1">Benzer S (November 1959). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC222769">"On the Topology of the Genetic Fine Structure"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>45</b> (11): <span class="nowrap">1607–</span>20. <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/1959PNAS...45.1607B">1959PNAS...45.1607B</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.45.11.1607">10.1073/pnas.45.11.1607</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/PMC222769">222769</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/16590553">16590553</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=On+the+Topology+of+the+Genetic+Fine+Structure&rft.volume=45&rft.issue=11&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1607-%3C%2Fspan%3E20&rft.date=1959-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC222769%23id-name%3DPMC&rft_id=info%3Apmid%2F16590553&rft_id=info%3Adoi%2F10.1073%2Fpnas.45.11.1607&rft_id=info%3Abibcode%2F1959PNAS...45.1607B&rft.aulast=Benzer&rft.aufirst=S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC222769&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-Min_1972-44"><span class="mw-cite-backlink"><b><a href="#cite_ref-Min_1972_44-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFMin_JouHaegemanYsebaertFiers1972" class="citation journal cs1">Min Jou W, Haegeman G, Ysebaert M, Fiers W (May 1972). 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(September 2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2716027">"The knockout mouse project"</a>. <i>Nature Genetics</i>. <b>36</b> (9): <span class="nowrap">921–</span>4. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fng0904-921">10.1038/ng0904-921</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/PMC2716027">2716027</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/15340423">15340423</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Genetics&rft.atitle=The+knockout+mouse+project&rft.volume=36&rft.issue=9&rft.pages=%3Cspan+class%3D%22nowrap%22%3E921-%3C%2Fspan%3E4&rft.date=2004-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2716027%23id-name%3DPMC&rft_id=info%3Apmid%2F15340423&rft_id=info%3Adoi%2F10.1038%2Fng0904-921&rft.aulast=Austin&rft.aufirst=CP&rft.au=Battey%2C+JF&rft.au=Bradley%2C+A&rft.au=Bucan%2C+M&rft.au=Capecchi%2C+M&rft.au=Collins%2C+FS&rft.au=Dove%2C+WF&rft.au=Duyk%2C+G&rft.au=Dymecki%2C+S&rft.au=Eppig%2C+JT&rft.au=Grieder%2C+FB&rft.au=Heintz%2C+N&rft.au=Hicks%2C+G&rft.au=Insel%2C+TR&rft.au=Joyner%2C+A&rft.au=Koller%2C+BH&rft.au=Lloyd%2C+KC&rft.au=Magnuson%2C+T&rft.au=Moore%2C+MW&rft.au=Nagy%2C+A&rft.au=Pollock%2C+JD&rft.au=Roses%2C+AD&rft.au=Sands%2C+AT&rft.au=Seed%2C+B&rft.au=Skarnes%2C+WC&rft.au=Snoddy%2C+J&rft.au=Soriano%2C+P&rft.au=Stewart%2C+DJ&rft.au=Stewart%2C+F&rft.au=Stillman%2C+B&rft.au=Varmus%2C+H&rft.au=Varticovski%2C+L&rft.au=Verma%2C+IM&rft.au=Vogt%2C+TF&rft.au=von+Melchner%2C+H&rft.au=Witkowski%2C+J&rft.au=Woychik%2C+RP&rft.au=Wurst%2C+W&rft.au=Yancopoulos%2C+GD&rft.au=Young%2C+SG&rft.au=Zambrowicz%2C+B&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2716027&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-123"><span class="mw-cite-backlink"><b><a href="#cite_ref-123">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGuanYeYangGao2010" class="citation journal cs1">Guan C, Ye C, Yang X, Gao J (February 2010). "A review of current large-scale mouse knockout efforts". <i>Genesis</i>. <b>48</b> (2): <span class="nowrap">73–</span>85. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fdvg.20594">10.1002/dvg.20594</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/20095055">20095055</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:34470273">34470273</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genesis&rft.atitle=A+review+of+current+large-scale+mouse+knockout+efforts&rft.volume=48&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E73-%3C%2Fspan%3E85&rft.date=2010-02&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A34470273%23id-name%3DS2CID&rft_id=info%3Apmid%2F20095055&rft_id=info%3Adoi%2F10.1002%2Fdvg.20594&rft.aulast=Guan&rft.aufirst=C&rft.au=Ye%2C+C&rft.au=Yang%2C+X&rft.au=Gao%2C+J&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> <li id="cite_note-Deng_2007-124"><span class="mw-cite-backlink"><b><a href="#cite_ref-Deng_2007_124-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFDeng2007" class="citation journal cs1">Deng C (October 2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2043165">"In celebration of Dr. Mario R. Capecchi's Nobel Prize"</a>. <i>International Journal of Biological Sciences</i>. <b>3</b> (7): <span class="nowrap">417–</span>9. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.7150%2Fijbs.3.417">10.7150/ijbs.3.417</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/PMC2043165">2043165</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/17998949">17998949</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=International+Journal+of+Biological+Sciences&rft.atitle=In+celebration+of+Dr.+Mario+R.+Capecchi%27s+Nobel+Prize&rft.volume=3&rft.issue=7&rft.pages=%3Cspan+class%3D%22nowrap%22%3E417-%3C%2Fspan%3E9&rft.date=2007-10&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2043165%23id-name%3DPMC&rft_id=info%3Apmid%2F17998949&rft_id=info%3Adoi%2F10.7150%2Fijbs.3.417&rft.aulast=Deng&rft.aufirst=C&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2043165&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading3"><h3 id="Sources">Sources</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene&action=edit&section=30" title="Edit section: Sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <dl><dt>Main textbook</dt></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFAlbertsJohnsonLewisRaff2002" class="citation book cs1"><a href="/wiki/Bruce_Alberts" title="Bruce Alberts">Alberts B</a>, Johnson A, <a href="/wiki/Julian_Lewis_(biologist)" title="Julian Lewis (biologist)">Lewis J</a>, <a href="/wiki/Martin_Raff" title="Martin Raff">Raff M</a>, Roberts K, <a href="/wiki/Peter_Walter" title="Peter Walter">Walter P</a> (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21054/"><i>Molecular Biology of the Cell</i></a> (Fourth ed.). New York: Garland Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-3218-3" title="Special:BookSources/978-0-8153-3218-3"><bdi>978-0-8153-3218-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+Biology+of+the+Cell&rft.place=New+York&rft.edition=Fourth&rft.pub=Garland+Science&rft.date=2002&rft.isbn=978-0-8153-3218-3&rft.aulast=Alberts&rft.aufirst=B&rft.au=Johnson%2C+A&rft.au=Lewis%2C+J&rft.au=Raff%2C+M&rft.au=Roberts%2C+K&rft.au=Walter%2C+P&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2FNBK21054%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span> – A molecular biology textbook available free online through NCBI Bookshelf.<span style="position:relative; top: -2px;"><span typeof="mw:File"><a href="/wiki/Open_access" title="open access publication – free to read"><img alt="Open access icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/Open_Access_logo_PLoS_transparent.svg/20px-Open_Access_logo_PLoS_transparent.svg.png" decoding="async" width="9" height="14" class="mw-file-element" data-file-width="640" data-file-height="1000" /></a></span></span></li></ul> <style data-mw-deduplicate="TemplateStyles:r1214851843">.mw-parser-output .hidden-begin{box-sizing:border-box;width:100%;padding:5px;border:none;font-size:95%}.mw-parser-output .hidden-title{font-weight:bold;line-height:1.6;text-align:left}.mw-parser-output .hidden-content{text-align:left}@media all and (max-width:500px){.mw-parser-output .hidden-begin{width:auto!important;clear:none!important;float:none!important}}</style><div class="hidden-begin mw-collapsible mw-collapsed" style="border:width: 1px; border-color:#999999;; width: 95%"><div class="hidden-title skin-nightmode-reset-color" style="background: #DBE9F4; padding-left:4px;">Referenced chapters of <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21054/"><i>Molecular Biology of the Cell</i></a></div><div class="hidden-content mw-collapsible-content" style=""> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21052/">Glossary</a></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26864/">Ch 1: Cells and genomes</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26864/">1.1: The Universal Features of Cells on Earth</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21068/">Ch 2: Cell Chemistry and Biosynthesis</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26883/">2.1: The Chemical Components of a Cell</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26830/">Ch 3: Proteins</a></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21074/">Ch 4: DNA and Chromosomes</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26821/">4.1: The Structure and Function of DNA</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26834/">4.2: Chromosomal DNA and Its Packaging in the Chromatin Fiber</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21064/">Ch 5: DNA Replication, Repair, and Recombination</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26850/">5.2: DNA Replication Mechanisms</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26879/">5.4: DNA Repair</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26898/">5.5: General Recombination</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21050/">Ch 6: How Cells Read the Genome: From DNA to Protein</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26887/">6.1: DNA to RNA</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26829/">6.2: RNA to Protein</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21057/">Ch 7: Control of Gene Expression</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26885/">7.1: An Overview of Gene Control</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26806/">7.2: DNA-Binding Motifs in Gene Regulatory Proteins</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26872/">7.3: How Genetic Switches Work</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26890/">7.5: Posttranscriptional Controls</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26836/">7.6: How Genomes Evolve</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21063/">Ch 14: Energy Conversion: Mitochondria and Chloroplasts</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26924/">14.4: The Genetic Systems of Mitochondria and Plastids</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21065/">Ch 18: The Mechanics of Cell Division</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26931/">18.1: An Overview of M Phase</a></dd> <dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26934/">18.2: Mitosis</a></dd></dl></dd></dl> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21049/">Ch 20: Germ Cells and Fertilization</a> <dl><dd><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK26840/">20.2: Meiosis</a></dd></dl></dd></dl> </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=Gene&action=edit&section=31" 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" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWatsonBakerBellGann2013" class="citation book cs1"><a href="/wiki/James_Watson" title="James Watson">Watson JD</a>, <a href="/wiki/Tania_A._Baker" title="Tania A. Baker">Baker TA</a>, Bell SP, Gann A, <a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Levine M</a>, Losick R (2013). <i>Molecular Biology of the Gene</i> (7th ed.). Benjamin Cummings. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-321-90537-6" title="Special:BookSources/978-0-321-90537-6"><bdi>978-0-321-90537-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+Biology+of+the+Gene&rft.edition=7th&rft.pub=Benjamin+Cummings&rft.date=2013&rft.isbn=978-0-321-90537-6&rft.aulast=Watson&rft.aufirst=JD&rft.au=Baker%2C+TA&rft.au=Bell%2C+SP&rft.au=Gann%2C+A&rft.au=Levine%2C+M&rft.au=Losick%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFDawkins1990" class="citation book cs1"><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Dawkins R</a> (1990). <a href="/wiki/The_Selfish_Gene" title="The Selfish Gene"><i>The Selfish Gene</i></a>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-286092-7" title="Special:BookSources/978-0-19-286092-7"><bdi>978-0-19-286092-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Selfish+Gene&rft.pub=Oxford+University+Press&rft.date=1990&rft.isbn=978-0-19-286092-7&rft.aulast=Dawkins&rft.aufirst=R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFRidley1999" class="citation book cs1"><a href="/wiki/Matt_Ridley" title="Matt Ridley">Ridley M</a> (1999). <a href="/wiki/Genome:_The_Autobiography_of_a_Species_in_23_Chapters" class="mw-redirect" title="Genome: The Autobiography of a Species in 23 Chapters"><i>Genome: The Autobiography of a Species in 23 Chapters</i></a>. Fourth Estate. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-00-763573-3" title="Special:BookSources/978-0-00-763573-3"><bdi>978-0-00-763573-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genome%3A+The+Autobiography+of+a+Species+in+23+Chapters&rft.pub=Fourth+Estate&rft.date=1999&rft.isbn=978-0-00-763573-3&rft.aulast=Ridley&rft.aufirst=M&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFBrown2002" class="citation book cs1">Brown T (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK21128"><i>Genomes</i></a> (2nd ed.). New York: Wiley-Liss. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-471-25046-3" title="Special:BookSources/978-0-471-25046-3"><bdi>978-0-471-25046-3</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/20821850">20821850</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20201106150314/https://www.ncbi.nlm.nih.gov/books/NBK21128/">Archived</a> from the original on 6 November 2020<span class="reference-accessdate">. Retrieved <span class="nowrap">31 August</span> 2017</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genomes&rft.place=New+York&rft.edition=2nd&rft.pub=Wiley-Liss&rft.date=2002&rft_id=info%3Apmid%2F20821850&rft.isbn=978-0-471-25046-3&rft.aulast=Brown&rft.aufirst=T&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2FNBK21128&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene" 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=Gene&action=edit&section=32" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1184024115" /><div class="div-col" style="column-width: 27em;"> <ul><li><a rel="nofollow" class="external text" href="https://ctdbase.org/">Comparative Toxicogenomics Database</a></li> <li><a rel="nofollow" class="external text" href="https://www.dnaftb.org/">DNA From The Beginning – a primer on genes and DNA</a></li> <li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/gene">Gene – a searchable database of genes</a></li> <li><a rel="nofollow" class="external text" href="https://www.mdpi.com/journal/genes"><i>Genes</i></a> – an Open Access journal</li> <li><a rel="nofollow" class="external text" href="http://idconverter.bioinfo.cnio.es/">IDconverter – converts gene IDs between public databases</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200929092205/http://idconverter.bioinfo.cnio.es/">Archived</a> 29 September 2020 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20051017154959/http://www.ihop-net.org/UniPub/iHOP/">iHOP – Information Hyperlinked over Proteins</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20110720065115/http://tagc.univ-mrs.fr/tbrowser/">TranscriptomeBrowser – Gene expression profile analysis</a></li> <li><a rel="nofollow" class="external text" href="https://archive.today/20121221003541/http://www.jcvi.org/pn-utility">The Protein Naming Utility, a database to identify and correct deficient gene names</a></li> <li><a rel="nofollow" class="external text" href="https://www.mousephenotype.org/">IMPC (International Mouse Phenotyping Consortium)</a> – Encyclopedia of mammalian gene function</li> <li><a rel="nofollow" class="external text" href="https://www.globalgenes.org/">Global Genes Project</a> – Leading non-profit organization supporting people living with genetic diseases</li> <li><a rel="nofollow" class="external text" href="https://www.nature.com/collections/aghcdefffg">Encode threads explorer</a>, <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i> <ul><li><a rel="nofollow" class="external text" href="https://www.nature.com/articles/nature28172">Characterization of intergenic regions and gene definition</a>, <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i></li></ul></li></ul> </div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output 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.navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Gene_expression217" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231" /><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression217" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction<br />to genetics</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a href="/wiki/DNA" title="DNA">DNA</a> → <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a> → <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA / hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Translation_(biology)" title="Translation (biology)">Translation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /></div><div role="navigation" class="navbox" aria-labelledby="Self-replicating_organic_structures216" 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_structures216" 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 class="mw-selflink selflink">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a 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 class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319" /></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q7187#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata1693" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div 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mw-ui-icon-wikimedia-history mw-ui-icon-wikimedia-wikimedia-history"></span> <span></span> </a> <a href="#" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only mw-watchlink" id="ca-watchstar-sticky-header" tabindex="-1" data-event-name="watch-sticky-header"><span class="vector-icon mw-ui-icon-wikimedia-star mw-ui-icon-wikimedia-wikimedia-star"></span> <span></span> </a> <a href="#" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only" id="ca-edit-sticky-header" tabindex="-1" data-event-name="wikitext-edit-sticky-header"><span class="vector-icon mw-ui-icon-wikimedia-wikiText mw-ui-icon-wikimedia-wikimedia-wikiText"></span> <span></span> </a> <a href="#" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only" id="ca-ve-edit-sticky-header" tabindex="-1" data-event-name="ve-edit-sticky-header"><span class="vector-icon mw-ui-icon-wikimedia-edit mw-ui-icon-wikimedia-wikimedia-edit"></span> <span></span> </a> <a href="#" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only" id="ca-viewsource-sticky-header" tabindex="-1" data-event-name="ve-edit-protected-sticky-header"><span class="vector-icon mw-ui-icon-wikimedia-editLock mw-ui-icon-wikimedia-wikimedia-editLock"></span> <span></span> </a> </div> <div class="vector-sticky-header-buttons"> <button class="cdx-button cdx-button--weight-quiet mw-interlanguage-selector" id="p-lang-btn-sticky-header" tabindex="-1" data-event-name="ui.dropdown-p-lang-btn-sticky-header"><span class="vector-icon mw-ui-icon-wikimedia-language mw-ui-icon-wikimedia-wikimedia-language"></span> <span>117 languages</span> </button> <a href="#" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive" id="ca-addsection-sticky-header" tabindex="-1" data-event-name="addsection-sticky-header"><span class="vector-icon mw-ui-icon-speechBubbleAdd-progressive mw-ui-icon-wikimedia-speechBubbleAdd-progressive"></span> <span>Add topic</span> </a> </div> <div class="vector-sticky-header-icon-end"> <div class="vector-user-links"> </div> </div> </div> </div> </div> <div class="mw-portlet mw-portlet-dock-bottom emptyPortlet" id="p-dock-bottom"> <ul> </ul> </div> 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