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processes</span> </div> </a> <button aria-controls="toc-Four_processes-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 Four processes subsection</span> </button> <ul id="toc-Four_processes-sublist" class="vector-toc-list"> <li id="toc-Selection" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Selection"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Selection</span> </div> </a> <ul id="toc-Selection-sublist" class="vector-toc-list"> <li id="toc-Dominance" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Dominance"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>Dominance</span> </div> </a> <ul id="toc-Dominance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Epistasis" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Epistasis"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Epistasis</span> </div> </a> <ul id="toc-Epistasis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mutation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mutation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Mutation</span> </div> </a> <ul id="toc-Mutation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_drift" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_drift"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Genetic drift</span> </div> </a> <ul id="toc-Genetic_drift-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gene_flow" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Gene_flow"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Gene flow</span> </div> </a> <ul id="toc-Gene_flow-sublist" class="vector-toc-list"> <li id="toc-Horizontal_gene_transfer" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Horizontal_gene_transfer"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4.1</span> <span>Horizontal gene transfer</span> </div> </a> <ul id="toc-Horizontal_gene_transfer-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Linkage" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Linkage"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Linkage</span> </div> </a> <ul id="toc-Linkage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Applications" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Applications"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Applications</span> </div> </a> <button aria-controls="toc-Applications-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Applications subsection</span> </button> <ul id="toc-Applications-sublist" class="vector-toc-list"> <li id="toc-Explaining_levels_of_genetic_variation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Explaining_levels_of_genetic_variation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Explaining levels of genetic variation</span> </div> </a> <ul id="toc-Explaining_levels_of_genetic_variation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Detecting_selection" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Detecting_selection"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Detecting selection</span> </div> </a> <ul id="toc-Detecting_selection-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Demographic_inference" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Demographic_inference"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Demographic inference</span> </div> </a> <ul id="toc-Demographic_inference-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_genetic_systems" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution_of_genetic_systems"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Evolution of genetic systems</span> </div> </a> <ul id="toc-Evolution_of_genetic_systems-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</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 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</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" > <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">Population genetics</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 53 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-53" 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">53 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%88%D8%B1%D8%A7%D8%AB%D9%8A%D8%A7%D8%AA_%D8%B3%D9%83%D8%A7%D9%86%D9%8A%D8%A9" title="وراثيات سكانية – Arabic" lang="ar" hreflang="ar" data-title="وراثيات سكانية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Populyasiya_genetikas%C4%B1" title="Populyasiya genetikası – Azerbaijani" lang="az" hreflang="az" data-title="Populyasiya genetikası" 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%9C%E0%A6%A8%E0%A6%B8%E0%A6%82%E0%A6%96%E0%A7%8D%E0%A6%AF%E0%A6%BE_%E0%A6%AC%E0%A6%82%E0%A6%B6%E0%A6%BE%E0%A6%A3%E0%A7%81%E0%A6%AC%E0%A6%BF%E0%A6%9C%E0%A7%8D%E0%A6%9E%E0%A6%BE%E0%A6%A8" title="জনসংখ্যা বংশাণুবিজ্ঞান – Bangla" lang="bn" hreflang="bn" data-title="জনসংখ্যা বংশাণুবিজ্ঞান" 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/K%C3%BBn-th%C3%A9_%C3%BBi-tho%C3%A2n-ha%CC%8Dk" title="Kûn-thé ûi-thoân-ha̍k – Minnan" lang="nan" hreflang="nan" data-title="Kûn-thé ûi-thoân-ha̍k" 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-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9F%D0%B0%D0%BF%D1%83%D0%BB%D1%8F%D1%86%D1%8B%D0%B9%D0%BD%D0%B0%D1%8F_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D1%8B%D0%BA%D0%B0" title="Папуляцыйная генетыка – Belarusian" lang="be" hreflang="be" data-title="Папуляцыйная генетыка" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BF%D1%83%D0%BB%D0%B0%D1%86%D0%B8%D0%BE%D0%BD%D0%BD%D0%B0_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Популационна генетика – Bulgarian" lang="bg" hreflang="bg" data-title="Популационна генетика" 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/Populacijska_genetika" title="Populacijska genetika – Bosnian" lang="bs" hreflang="bs" data-title="Populacijska genetika" 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%C3%A8tica_de_poblacions" title="Genètica de poblacions – Catalan" lang="ca" hreflang="ca" data-title="Genètica de poblacions" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Popula%C4%8Dn%C3%AD_genetika" title="Populační genetika – Czech" lang="cs" hreflang="cs" data-title="Populační genetika" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Populationsgenetik" title="Populationsgenetik – German" lang="de" hreflang="de" data-title="Populationsgenetik" 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/Populatsioonigeneetika" title="Populatsioonigeneetika – Estonian" lang="et" hreflang="et" data-title="Populatsioonigeneetika" 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%B5%CE%BD%CE%B5%CF%84%CE%B9%CE%BA%CE%AE_%CF%84%CF%89%CE%BD_%CF%80%CE%BB%CE%B7%CE%B8%CF%85%CF%83%CE%BC%CF%8E%CE%BD" title="Γενετική των πληθυσμών – Greek" lang="el" hreflang="el" data-title="Γενετική των πληθυσμών" 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%C3%A9tica_de_poblaciones" title="Genética de poblaciones – Spanish" lang="es" hreflang="es" data-title="Genética de poblaciones" 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/Populacigenetiko" title="Populacigenetiko – Esperanto" lang="eo" hreflang="eo" data-title="Populacigenetiko" 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/Populazioen_genetika" title="Populazioen genetika – Basque" lang="eu" hreflang="eu" data-title="Populazioen genetika" 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%D8%AA%DB%8C%DA%A9_%D8%AC%D9%85%D8%B9%DB%8C%D8%AA" title="ژنتیک جمعیت – Persian" lang="fa" hreflang="fa" data-title="ژنتیک جمعیت" 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%A9n%C3%A9tique_des_populations" title="Génétique des populations – French" lang="fr" hreflang="fr" data-title="Génétique des populations" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Xen%C3%A9tica_de_poboaci%C3%B3ns" title="Xenética de poboacións – Galician" lang="gl" hreflang="gl" data-title="Xenética de poboacións" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%A7%91%EB%8B%A8%EC%9C%A0%EC%A0%84%ED%95%99" title="집단유전학 – Korean" lang="ko" hreflang="ko" data-title="집단유전학" 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/%D5%8A%D5%B8%D5%BA%D5%B8%D6%82%D5%AC%D5%B5%D5%A1%D6%81%D5%AB%D5%B8%D5%B6_%D5%A3%D5%A5%D5%B6%D5%A5%D5%BF%D5%AB%D5%AF%D5%A1" title="Պոպուլյացիոն գենետիկա – Armenian" lang="hy" hreflang="hy" data-title="Պոպուլյացիոն գենետիկա" 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%A4%A8%E0%A4%B8%E0%A4%82%E0%A4%96%E0%A5%8D%E0%A4%AF%E0%A4%BE_%E0%A4%86%E0%A4%A8%E0%A5%81%E0%A4%B5%E0%A4%BE%E0%A4%82%E0%A4%B6%E0%A4%BF%E0%A4%95%E0%A5%80" title="जनसंख्या आनुवांशिकी – Hindi" lang="hi" hreflang="hi" data-title="जनसंख्या आनुवांशिकी" 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/Populacijska_genetika" title="Populacijska genetika – Croatian" lang="hr" hreflang="hr" data-title="Populacijska genetika" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Genetika_populasi" title="Genetika populasi – Indonesian" lang="id" hreflang="id" data-title="Genetika populasi" 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-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Stofnerf%C3%B0afr%C3%A6%C3%B0i" title="Stofnerfðafræði – Icelandic" lang="is" hreflang="is" data-title="Stofnerfðafræði" 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/Genetica_delle_popolazioni" title="Genetica delle popolazioni – Italian" lang="it" hreflang="it" data-title="Genetica delle popolazioni" 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%A0%D7%98%D7%99%D7%A7%D7%94_%D7%A9%D7%9C_%D7%90%D7%95%D7%9B%D7%9C%D7%95%D7%A1%D7%99%D7%95%D7%AA" title="גנטיקה של אוכלוסיות – Hebrew" lang="he" hreflang="he" data-title="גנטיקה של אוכלוסיות" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Genetica_popularis" title="Genetica popularis – Latin" lang="la" hreflang="la" data-title="Genetica popularis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Popul%C3%A1ci%C3%B3genetika" title="Populációgenetika – Hungarian" lang="hu" hreflang="hu" data-title="Populációgenetika" 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%9F%D0%BE%D0%BF%D1%83%D0%BB%D0%B0%D1%86%D0%B8%D0%BE%D0%BD%D0%B0_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Популациона генетика – Macedonian" lang="mk" hreflang="mk" data-title="Популациона генетика" 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%B4%A8%E0%B4%B8%E0%B4%82%E0%B4%96%E0%B5%8D%E0%B4%AF%E0%B4%BE%E0%B4%9C%E0%B4%A8%E0%B4%BF%E0%B4%A4%E0%B4%95%E0%B4%B6%E0%B4%BE%E0%B4%B8%E0%B5%8D%E0%B4%A4%E0%B5%8D%E0%B4%B0%E0%B4%82" title="ജനസംഖ്യാജനിതകശാസ്ത്രം – Malayalam" lang="ml" hreflang="ml" data-title="ജനസംഖ്യാജനിതകശാസ്ത്രം" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Genetik_populasi" title="Genetik populasi – Malay" lang="ms" hreflang="ms" data-title="Genetik populasi" 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-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BF%D1%83%D0%BB%D1%8F%D1%86%D0%B8%D0%B9%D0%BD_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA" title="Популяцийн генетик – Mongolian" lang="mn" hreflang="mn" data-title="Популяцийн генетик" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Populatiegenetica" title="Populatiegenetica – Dutch" lang="nl" hreflang="nl" data-title="Populatiegenetica" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E9%9B%86%E5%9B%A3%E9%81%BA%E4%BC%9D%E5%AD%A6" title="集団遺伝学 – Japanese" lang="ja" hreflang="ja" data-title="集団遺伝学" 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-nov mw-list-item"><a href="https://nov.wikipedia.org/wiki/Populational_genetike" title="Populational genetike – Novial" lang="nov" hreflang="nov" data-title="Populational genetike" data-language-autonym="Novial" data-language-local-name="Novial" class="interlanguage-link-target"><span>Novial</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Genetica_de_las_populacions" title="Genetica de las populacions – Occitan" lang="oc" hreflang="oc" data-title="Genetica de las populacions" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AF_%D9%88%DA%AB%DA%93%D9%88_%DB%8C%D8%A7_%DA%89%D9%84%D9%88_%D8%AC%D9%86%DB%8C%D9%BC%DB%8C%DA%A9" title="د وګړو یا ډلو جنیټیک – Pashto" lang="ps" hreflang="ps" data-title="د وګړو یا ډلو جنیټیک" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Genetyka_populacyjna" title="Genetyka populacyjna – Polish" lang="pl" hreflang="pl" data-title="Genetyka populacyjna" 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/Gen%C3%A9tica_populacional" title="Genética populacional – Portuguese" lang="pt" hreflang="pt" data-title="Genética populacional" 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/Genetica_popula%C8%9Biilor" title="Genetica populațiilor – Romanian" lang="ro" hreflang="ro" data-title="Genetica populațiilor" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BF%D1%83%D0%BB%D1%8F%D1%86%D0%B8%D0%BE%D0%BD%D0%BD%D0%B0%D1%8F_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Популяционная генетика – Russian" lang="ru" hreflang="ru" data-title="Популяционная генетика" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Population_genetics" title="Population genetics – Simple English" lang="en-simple" hreflang="en-simple" data-title="Population genetics" 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-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BF%D1%83%D0%BB%D0%B0%D1%86%D0%B8%D0%BE%D0%BD%D0%B0_%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0" title="Популациона генетика – Serbian" lang="sr" hreflang="sr" data-title="Популациона генетика" 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/Populaciona_genetika" title="Populaciona genetika – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Populaciona genetika" 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/Populaatiogenetiikka" title="Populaatiogenetiikka – Finnish" lang="fi" hreflang="fi" data-title="Populaatiogenetiikka" 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/Populationsgenetik" title="Populationsgenetik – Swedish" lang="sv" hreflang="sv" data-title="Populationsgenetik" 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/Henetikang_pampopulasyon" title="Henetikang pampopulasyon – Tagalog" lang="tl" hreflang="tl" data-title="Henetikang pampopulasyon" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%9E%E0%B8%B1%E0%B8%99%E0%B8%98%E0%B8%B8%E0%B8%A8%E0%B8%B2%E0%B8%AA%E0%B8%95%E0%B8%A3%E0%B9%8C%E0%B8%9B%E0%B8%A3%E0%B8%B0%E0%B8%8A%E0%B8%B2%E0%B8%81%E0%B8%A3" title="พันธุศาสตร์ประชากร – Thai" lang="th" hreflang="th" data-title="พันธุศาสตร์ประชากร" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Pop%C3%BClasyon_geneti%C4%9Fi" title="Popülasyon genetiği – Turkish" lang="tr" hreflang="tr" data-title="Popülasyon genetiği" 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%D0%B5%D1%82%D0%B8%D0%BA%D0%B0_%D0%BF%D0%BE%D0%BF%D1%83%D0%BB%D1%8F%D1%86%D1%96%D0%B9" title="Генетика популяцій – Ukrainian" lang="uk" hreflang="uk" data-title="Генетика популяцій" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Di_truy%E1%BB%81n_h%E1%BB%8Dc_qu%E1%BA%A7n_th%E1%BB%83" title="Di truyền học quần thể – Vietnamese" lang="vi" hreflang="vi" data-title="Di truyền học quần thể" 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-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%BE%A4%E9%AB%94%E9%81%BA%E5%82%B3%E5%AD%B8" title="群體遺傳學 – 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rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><table class="sidebar sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><span class="notpageimage" typeof="mw:File"><a href="/wiki/File:Darwin%27s_finches_by_Gould.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/180px-Darwin%27s_finches_by_Gould.jpg" decoding="async" width="180" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/270px-Darwin%27s_finches_by_Gould.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/360px-Darwin%27s_finches_by_Gould.jpg 2x" data-file-width="1748" data-file-height="1319" /></a></span><div class="sidebar-caption"><a href="/wiki/Darwin%27s_finches" title="Darwin's finches">Darwin's finches</a> by <a href="/wiki/John_Gould" title="John Gould">John Gould</a></div></td></tr><tr><td class="sidebar-above" style="display:block; margin-bottom:0.55em; background-color: transparent;"> <div class="hlist"> <ul><li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li> <li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Evolution" title="Evolution">Main</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li></ul> <div class="hlist"> </div> <ul><li><a href="/wiki/Glossary_of_evolutionary_biology" class="mw-redirect" title="Glossary of evolutionary biology">Glossary</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence</a></li> <li><a href="/wiki/History_of_life" title="History of life">History</a></li></ul> </div></td></tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Processes and outcomes</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a class="mw-selflink selflink">Population genetics</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Diversity</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Co-operation</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Coextinction" title="Coextinction">Coextinction</a></li> <li><a href="/wiki/Contingency_(evolutionary_biology)" title="Contingency (evolutionary biology)">Contingency</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a> <ul><li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li> <li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Extinction_event" title="Extinction event">Extinction event</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li> <li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Darwin</a></li> <li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">Origin of Species</a></i></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evo-devo</a></li> <li><a href="/wiki/Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History of speciation</a></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="/wiki/Timeline_of_paleontology" title="Timeline of paleontology">timeline</a>)</li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li> <li><a href="/wiki/Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li> <li><a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li> <li><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li> <li><a href="/wiki/Experimental_evolution" title="Experimental evolution">Experimental evolution</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li> <li><a 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title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li> <li><a href="/wiki/Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li> <li><a href="/wiki/Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li> <li><a href="/wiki/Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="nowrap"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Tree_of_life.svg" class="mw-file-description"><img alt="icon" 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href="/wiki/Special:EditPage/Template:Evolutionary_biology" title="Special:EditPage/Template:Evolutionary biology"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1246091330"><table class="sidebar nomobile plainlist" style="width:17em"><tbody><tr><td class="sidebar-pretitle">Part of <a href="/wiki/Category:Genetic_genealogy" title="Category:Genetic genealogy">a series</a> on</td></tr><tr><th class="sidebar-title-with-pretitle" style="font-size:150%;"><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></th></tr><tr><th class="sidebar-heading" style="background:#ddd;padding-bottom:0.25em"> Concepts</th></tr><tr><td class="sidebar-content"> <ul><li><a class="mw-selflink selflink">Population genetics</a></li> <li><a href="/wiki/Haplogroup" title="Haplogroup">Haplogroup</a> / <a href="/wiki/Haplotype" title="Haplotype">Haplotype</a> / <a href="/wiki/Subclade" title="Subclade">Subclade</a></li> <li><a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">Most recent common ancestor</a></li> <li><div style="display:inline-block; padding:0.2em 0.4em; line-height:1.2em;"><a href="/wiki/Haplogroup" title="Haplogroup">DNA haplogroups</a><br />(<a href="/wiki/Human_mitochondrial_DNA_haplogroup" title="Human mitochondrial DNA haplogroup">Human mitochondrial</a>,<br /><a href="/wiki/Human_Y-chromosome_DNA_haplogroup" title="Human Y-chromosome DNA haplogroup">Human Y-chromosome</a>)</div></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li></ul></td> </tr><tr><th class="sidebar-heading" style="background:#ddd;padding-bottom:0.25em"> Related topics</th></tr><tr><td class="sidebar-content"> <ul><li><div style="display:inline-block; padding:0.2em 0.4em; line-height:1.2em;"><a href="/wiki/Y-chromosome_haplogroups_in_populations_of_the_world" class="mw-redirect" title="Y-chromosome haplogroups in populations of the world">Y-chromosome haplogroups by population</a></div></li> <li><a href="/wiki/Genealogical_DNA_test" title="Genealogical DNA test">Genealogical DNA test</a></li> <li><a href="/wiki/Surname_DNA_project" title="Surname DNA project">Surname DNA project</a></li> <li><a href="/wiki/Personal_genomics" title="Personal genomics">Personal genomics</a></li> <li><a href="/wiki/Genographic_Project" title="Genographic Project">Genographic Project</a></li> <li><a href="/wiki/International_Society_of_Genetic_Genealogy" title="International Society of Genetic Genealogy">ISOGG</a></li></ul></td> </tr><tr><td class="sidebar-navbar"><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:Genetic_genealogy" title="Template:Genetic genealogy"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetic_genealogy" title="Template talk:Genetic genealogy"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetic_genealogy" title="Special:EditPage/Template:Genetic genealogy"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p><b>Population genetics</b> is a subfield of <a href="/wiki/Genetics" title="Genetics">genetics</a> that deals with genetic differences within and among <a href="/wiki/Population" title="Population">populations</a>, and is a part of <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>. Studies in this branch of <a href="/wiki/Biology" title="Biology">biology</a> examine such phenomena as <a href="/wiki/Adaptation_(biology)" class="mw-redirect" title="Adaptation (biology)">adaptation</a>, <a href="/wiki/Speciation" title="Speciation">speciation</a>, and <a href="/wiki/Population_stratification" class="mw-redirect" title="Population stratification">population structure</a>.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Population genetics was a vital ingredient in the <a href="/wiki/Emergence" title="Emergence">emergence</a> of the <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern evolutionary synthesis</a>. Its primary founders were <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a>, <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> and <a href="/wiki/Ronald_Fisher" title="Ronald Fisher">Ronald Fisher</a>, who also laid the foundations for the related discipline of <a href="/wiki/Quantitative_genetics" title="Quantitative genetics">quantitative genetics</a>. Traditionally a highly mathematical discipline, modern population genetics encompasses theoretical, laboratory, and field work. Population genetic models are used both for <a href="/wiki/Statistical_inference" title="Statistical inference">statistical inference</a> from DNA sequence data and for proof/disproof of concept.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>What sets population genetics apart from newer, more phenotypic approaches to modelling evolution, such as <a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">evolutionary game theory</a> and <a href="/wiki/Evolutionary_invasion_analysis" title="Evolutionary invasion analysis">adaptive dynamics</a>, is its emphasis on such genetic phenomena as <a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">dominance</a>, <a href="/wiki/Epistasis" title="Epistasis">epistasis</a>, the degree to which <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> breaks <a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a>, and the random phenomena of <a href="/wiki/Mutation" title="Mutation">mutation</a> and <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>. This makes it appropriate for comparison to <a href="/wiki/Population_genomics" title="Population genomics">population genomics</a> data. </p> <meta property="mw:PageProp/toc" /> <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=Population_genetics&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Population genetics began as a reconciliation of <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a> and <a href="/wiki/Biostatistics" title="Biostatistics">biostatistics</a> models. <a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a> will only cause evolution if there is enough <a href="/wiki/Genetic_variation" title="Genetic variation">genetic variation</a> in a population. Before the discovery of <a href="/wiki/Mendelian_genetics" class="mw-redirect" title="Mendelian genetics">Mendelian genetics</a>, one common hypothesis was <a href="/wiki/Blending_inheritance" title="Blending inheritance">blending inheritance</a>. But with blending inheritance, genetic variance would be rapidly lost, making evolution by natural or sexual selection implausible. The <a href="/wiki/Hardy%E2%80%93Weinberg_principle" title="Hardy–Weinberg principle">Hardy–Weinberg principle</a> provides the solution to how variation is maintained in a population with Mendelian inheritance. According to this principle, the frequencies of alleles (variations in a gene) will remain constant in the absence of selection, mutation, migration and genetic drift.<sup id="cite_ref-Ewens_W.J._2004_3-0" class="reference"><a href="#cite_note-Ewens_W.J._2004-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:204px;max-width:204px"><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Biston.betularia.7200.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/200px-Biston.betularia.7200.jpg" decoding="async" width="200" height="119" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/300px-Biston.betularia.7200.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/400px-Biston.betularia.7200.jpg 2x" data-file-width="2798" data-file-height="1662" /></a></span></div><div class="thumbcaption">The typical white-bodied form of the <a href="/wiki/Peppered_moth" title="Peppered moth">peppered moth</a></div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Biston.betularia.f.carbonaria.7209.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/200px-Biston.betularia.f.carbonaria.7209.jpg" decoding="async" width="200" height="97" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/300px-Biston.betularia.f.carbonaria.7209.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/400px-Biston.betularia.f.carbonaria.7209.jpg 2x" data-file-width="2816" data-file-height="1366" /></a></span></div><div class="thumbcaption"><a href="/wiki/Industrial_melanism" title="Industrial melanism">Industrial melanism</a>: the black-bodied form of the peppered moth appeared in polluted areas.</div></div></div></div></div> <p>The next key step was the work of the British biologist and statistician <a href="/wiki/Ronald_Fisher" title="Ronald Fisher">Ronald Fisher</a>. In a series of papers starting in 1918 and culminating in his 1930 book <i><a href="/wiki/The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection">The Genetical Theory of Natural Selection</a></i>, Fisher showed that the continuous variation measured by the biometricians could be produced by the combined action of many discrete genes, and that natural selection could change allele frequencies in a population, resulting in evolution. In a series of papers beginning in 1924, another British geneticist, <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a>, worked out the mathematics of allele frequency change at a single gene <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">locus</a> under a broad range of conditions. Haldane also applied statistical analysis to real-world examples of natural selection, such as <a href="/wiki/Peppered_moth_evolution" title="Peppered moth evolution">peppered moth evolution</a> and <a href="/wiki/Industrial_melanism" title="Industrial melanism">industrial melanism</a>, and showed that <a href="/wiki/Selection_coefficient" title="Selection coefficient">selection coefficients</a> could be larger than Fisher assumed, leading to more rapid adaptive evolution as a camouflage strategy following increased pollution.<sup id="cite_ref-Bowler325-339_4-0" class="reference"><a href="#cite_note-Bowler325-339-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larson221-243_5-0" class="reference"><a href="#cite_note-Larson221-243-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The American biologist <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a>, who had a background in <a href="/wiki/Animal_breeding" title="Animal breeding">animal breeding</a> experiments, focused on combinations of interacting genes, and the effects of <a href="/wiki/Inbreeding" title="Inbreeding">inbreeding</a> on small, relatively isolated populations that exhibited genetic drift. In 1932 Wright introduced the concept of an <a href="/wiki/Fitness_landscape" title="Fitness landscape">adaptive landscape</a> and argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive peak, allowing natural selection to drive it towards different adaptive peaks.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2020)">citation needed</span></a></i>]</sup> </p><p>The work of Fisher, Haldane and Wright founded the discipline of population genetics. This integrated natural selection with Mendelian genetics, which was the critical first step in developing a unified theory of how evolution worked.<sup id="cite_ref-Bowler325-339_4-1" class="reference"><a href="#cite_note-Bowler325-339-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larson221-243_5-1" class="reference"><a href="#cite_note-Larson221-243-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">John Maynard Smith</a> was Haldane's pupil, whilst <a href="/wiki/W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a> was influenced by the writings of Fisher. The American <a href="/wiki/George_R._Price" title="George R. Price">George R. Price</a> worked with both Hamilton and Maynard Smith. American <a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a> and Japanese <a href="/wiki/Motoo_Kimura" title="Motoo Kimura">Motoo Kimura</a> were influenced by Wright and Haldane.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2020)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Modern_synthesis">Modern synthesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=2" title="Edit section: Modern synthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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">Main article: <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis (20th century)</a></div> <p>The mathematics of population genetics were originally developed as the beginning of the <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern synthesis</a>. Authors such as Beatty<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> have asserted that population genetics defines the core of the modern synthesis. For the first few decades of the 20th century, most field naturalists continued to believe that <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a> and <a href="/wiki/Orthogenesis" title="Orthogenesis">orthogenesis</a> provided the best explanation for the complexity they observed in the living world.<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> During the modern synthesis, these ideas were purged, and only evolutionary causes that could be expressed in the mathematical framework of population genetics were retained.<sup id="cite_ref-Provine88_8-0" class="reference"><a href="#cite_note-Provine88-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Consensus was reached as to which evolutionary factors might influence evolution, but not as to the relative importance of the various factors.<sup id="cite_ref-Provine88_8-1" class="reference"><a href="#cite_note-Provine88-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a>, a postdoctoral worker in <a href="/wiki/T._H._Morgan" class="mw-redirect" title="T. H. Morgan">T. H. Morgan</a>'s lab, had been influenced by the work on <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> by Russian geneticists such as <a href="/wiki/Sergei_Chetverikov" title="Sergei Chetverikov">Sergei Chetverikov</a>. He helped to bridge the divide between the foundations of <a href="/wiki/Microevolution" title="Microevolution">microevolution</a> developed by the population geneticists and the patterns of <a href="/wiki/Macroevolution" title="Macroevolution">macroevolution</a> observed by field biologists, with his 1937 book <i><a href="/wiki/Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species">Genetics and the Origin of Species</a></i>. Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the assumptions of the population geneticists, these populations had large amounts of genetic diversity, with marked differences between sub-populations. The book also took the highly mathematical work of the population geneticists and put it into a more accessible form. Many more biologists were influenced by population genetics via Dobzhansky than were able to read the highly mathematical works in the original.<sup id="cite_ref-Provine78_9-0" class="reference"><a href="#cite_note-Provine78-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>In Great Britain <a href="/wiki/E._B._Ford" title="E. B. Ford">E. B. Ford</a>, the pioneer of <a href="/wiki/Ecological_genetics" title="Ecological genetics">ecological genetics</a>,<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> continued throughout the 1930s and 1940s to empirically demonstrate the power of selection due to ecological factors including the ability to maintain genetic diversity through genetic <a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">polymorphisms</a> such as human <a href="/wiki/Blood_type" title="Blood type">blood types</a>. Ford's work, in collaboration with Fisher, contributed to a shift in emphasis during the modern synthesis towards natural selection as the dominant force.<sup id="cite_ref-Bowler325-339_4-2" class="reference"><a href="#cite_note-Bowler325-339-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larson221-243_5-2" class="reference"><a href="#cite_note-Larson221-243-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neutral_theory_and_origin-fixation_dynamics">Neutral theory and origin-fixation dynamics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=3" title="Edit section: Neutral theory and origin-fixation dynamics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The original, modern synthesis view of population genetics assumes that mutations provide ample raw material, and focuses only on the change in <a href="/wiki/Allele_frequency" title="Allele frequency">frequency of alleles</a> within <a href="/wiki/Population" title="Population">populations</a>.<sup id="cite_ref-McCandlish-2014_13-0" class="reference"><a href="#cite_note-McCandlish-2014-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The main processes influencing allele frequencies are <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>, <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>, <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> and recurrent <a href="/wiki/Mutation" title="Mutation">mutation</a>. Fisher and Wright had some fundamental disagreements about the relative roles of selection and drift.<sup id="cite_ref-Crow_2010_14-0" class="reference"><a href="#cite_note-Crow_2010-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The availability of molecular data on all genetic differences led to the <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of molecular evolution</a>. In this view, many mutations are deleterious and so never observed, and most of the remainder are neutral, i.e. are not under selection. With the fate of each neutral mutation left to chance (genetic drift), the direction of evolutionary change is driven by which mutations occur, and so cannot be captured by models of change in the frequency of (existing) alleles alone.<sup id="cite_ref-McCandlish-2014_13-1" class="reference"><a href="#cite_note-McCandlish-2014-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-CasillasBarbadilla2017_15-0" class="reference"><a href="#cite_note-CasillasBarbadilla2017-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>The origin-fixation view of population genetics generalizes this approach beyond strictly neutral mutations, and sees the rate at which a particular change happens as the product of the mutation rate and the <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixation probability</a>.<sup id="cite_ref-McCandlish-2014_13-2" class="reference"><a href="#cite_note-McCandlish-2014-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Four_processes">Four processes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=4" title="Edit section: Four processes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Selection">Selection</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=5" title="Edit section: Selection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a>, which includes <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>, is the fact that some <a href="/wiki/Trait_(biology)" class="mw-redirect" title="Trait (biology)">traits</a> make it more likely for an <a href="/wiki/Organism" title="Organism">organism</a> to survive and <a href="/wiki/Reproduction" title="Reproduction">reproduce</a>. Population genetics describes natural selection by defining <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> as a <a href="/wiki/Propensity_probability" title="Propensity probability">propensity or probability</a> of survival and reproduction in a particular environment. The fitness is normally given by the symbol <b>w</b>=1-<b>s</b> where <b>s</b> is the <a href="/wiki/Selection_coefficient" title="Selection coefficient">selection coefficient</a>. Natural selection acts on <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a>, so population genetic models assume relatively simple relationships to predict the phenotype and hence fitness from the <a href="/wiki/Allele" title="Allele">allele</a> at one or a small number of loci. In this way, natural selection converts differences in the fitness of individuals with different phenotypes into changes in allele frequency in a population over successive generations.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>Before the advent of population genetics, many biologists doubted that small differences in fitness were sufficient to make a large difference to evolution.<sup id="cite_ref-Provine78_9-1" class="reference"><a href="#cite_note-Provine78-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Population geneticists addressed this concern in part by comparing selection to <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>. Selection can overcome genetic drift when <b>s</b> is greater than 1 divided by the <a href="/wiki/Effective_population_size" title="Effective population size">effective population size</a>. When this criterion is met, the probability that a new advantageous mutant becomes <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixed</a> is approximately equal to <b>2s</b>.<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><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> The time until fixation of such an allele is approximately <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle (2log(sN)+\gamma )/s}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mo stretchy="false">(</mo> <mn>2</mn> <mi>l</mi> <mi>o</mi> <mi>g</mi> <mo stretchy="false">(</mo> <mi>s</mi> <mi>N</mi> <mo stretchy="false">)</mo> <mo>+</mo> <mi>γ</mi> <mo stretchy="false">)</mo> <mrow class="MJX-TeXAtom-ORD"> <mo>/</mo> </mrow> <mi>s</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle (2log(sN)+\gamma )/s}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/83cd931ec80a70830d1ed5d0ea331a084ab87bf6" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:17.228ex; height:2.843ex;" alt="{\displaystyle (2log(sN)+\gamma )/s}"></span>.<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> </p> <div class="mw-heading mw-heading4"><h4 id="Dominance">Dominance</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=6" title="Edit section: Dominance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a> means that the phenotypic and/or fitness effect of one allele at a locus depends on which allele is present in the second copy for that locus. Consider three genotypes at one locus, with the following fitness values<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> </p> <table class="wikitable"> <tbody><tr> <td>Genotype: </td> <td>A<sub>1</sub>A<sub>1</sub> </td> <td>A<sub>1</sub>A<sub>2</sub> </td> <td>A<sub>2</sub>A<sub>2</sub> </td></tr> <tr> <td>Relative fitness: </td> <td>1 </td> <td>1-hs </td> <td>1-s </td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox" style="width:452px; ; border-collapse:collapse;"><tbody><tr><th colspan="2" class="infobox-above" style="font-size:100%; background:#FFCC66; box-shadow: inset 3px 3px 0 #fafafa, inset -3px -3px 0 #fafafa;; padding:5px;">Population genetics glossary</th></tr><tr><td colspan="2" class="infobox-full-data" style="text-align:left; padding-left:0;"> <ul><li><b><a href="/wiki/Species" title="Species">species</a></b> – a group of closely related organisms which, if sexual, are capable of interbreeding and producing fertile offspring</li> <li><b><a href="/wiki/Population" title="Population">population</a></b> – the set of individuals of a particular species in a given area</li> <li><b><a href="/wiki/Gene_pool" title="Gene pool">gene pool</a></b> – the collective genetic information contained within a population of sexually reproducing organisms; ignores <a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a></li> <li><b><a href="/wiki/Allele_frequency" title="Allele frequency">allele frequency</a></b> – the frequency or proportion of a particular allele of a gene within a population</li></ul> </td></tr></tbody></table> <p>s is the <a href="/wiki/Selection_coefficient" title="Selection coefficient">selection coefficient</a> and h is the dominance coefficient. The value of h yields the following information: </p> <table class="wikitable"> <tbody><tr> <td>h=0 </td> <td>A<sub>1</sub> dominant, A<sub>2</sub> recessive </td></tr> <tr> <td>h=1 </td> <td>A<sub>2</sub> dominant, A<sub>1</sub> recessive </td></tr> <tr> <td>0<h<1 </td> <td>incomplete dominance </td></tr> <tr> <td>h<0 </td> <td><a href="/wiki/Overdominance" title="Overdominance">overdominance</a> </td></tr> <tr> <td>h>1 </td> <td><a href="/wiki/Underdominance" title="Underdominance">Underdominance</a> </td></tr></tbody></table> <div class="mw-heading mw-heading4"><h4 id="Epistasis">Epistasis</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=7" title="Edit section: Epistasis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Synergistic_versus_antagonistic_epistasis.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/200px-Synergistic_versus_antagonistic_epistasis.svg.png" decoding="async" width="200" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/300px-Synergistic_versus_antagonistic_epistasis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/400px-Synergistic_versus_antagonistic_epistasis.svg.png 2x" data-file-width="724" data-file-height="570" /></a><figcaption>The <a href="/wiki/Logarithm" title="Logarithm">logarithm</a> of fitness as a function of the number of deleterious mutations. Synergistic epistasis is represented by the red line - each subsequent deleterious mutation has a larger proportionate effect on the organism's fitness. Antagonistic epistasis is in blue. The black line shows the non-epistatic case, where fitness is the <a href="/wiki/Product_(mathematics)" title="Product (mathematics)">product</a> of the contributions from each of its loci.</figcaption></figure> <p><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a> means that the phenotypic and/or fitness effect of an allele at one locus depends on which alleles are present at other loci. Selection does not act on a single locus, but on a phenotype that arises through development from a complete genotype.<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> However, many population genetics models of sexual species are "single locus" models, where the fitness of an individual is calculated as the <a href="/wiki/Product_(mathematics)" title="Product (mathematics)">product</a> of the contributions from each of its loci—effectively assuming no epistasis. </p><p>In fact, the <a href="/wiki/Fitness_landscape" title="Fitness landscape">genotype to fitness landscape</a> is more complex. Population genetics must either model this complexity in detail, or capture it by some simpler average rule. Empirically, beneficial mutations tend to have a smaller fitness benefit when added to a genetic background that already has high fitness: this is known as diminishing returns epistasis.<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> When deleterious mutations also have a smaller fitness effect on high fitness backgrounds, this is known as "synergistic epistasis". However, the effect of deleterious mutations tends on average to be very close to multiplicative, or can even show the opposite pattern, known as "antagonistic epistasis".<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> </p><p>Synergistic epistasis is central to some theories of the purging of <a href="/wiki/Genetic_load" title="Genetic load">mutation load</a><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> and to the <a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">evolution of sexual reproduction</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Mutation">Mutation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=8" title="Edit section: Mutation"><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/Mutation" title="Mutation">Mutation</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Drosophila_melanogaster_-_side_(aka).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/240px-Drosophila_melanogaster_-_side_%28aka%29.jpg" decoding="async" width="240" height="186" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/360px-Drosophila_melanogaster_-_side_%28aka%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Drosophila_melanogaster_-_side_%28aka%29.jpg/480px-Drosophila_melanogaster_-_side_%28aka%29.jpg 2x" data-file-width="546" data-file-height="424" /></a><figcaption><i>Drosophila melanogaster</i></figcaption></figure> <p>The genetic process of <a href="/wiki/Mutation" title="Mutation">mutation</a> takes place within an individual, resulting in heritable changes to the genetic material. This process is often characterized by a description of the starting and ending states, or the kind of change that has happened at the level of DNA (e.g,. a T-to-C mutation, a 1-bp deletion), of genes or proteins (e.g., a null mutation, a loss-of-function mutation), or at a higher phenotypic level (e.g., red-eye mutation). Single-nucleotide changes are frequently the most common type of mutation, but many other types of <a href="/wiki/Mutation" title="Mutation">mutation</a> are possible, and they occur at widely varying rates that may show systematic asymmetries or biases (<a href="/wiki/Mutation_bias" title="Mutation bias">mutation bias</a>). </p><p>Mutations can involve large sections of DNA becoming <a href="/wiki/Gene_duplication" title="Gene duplication">duplicated</a>, usually through <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>.<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> This leads to <a href="/wiki/Copy-number_variation" class="mw-redirect" title="Copy-number variation">copy-number variation</a> within a population. Duplications are a major source of raw material for evolving new genes.<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> Other types of mutation occasionally create new genes from previously noncoding DNA.<sup id="cite_ref-26" class="reference"><a href="#cite_note-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> </p><p>In the <a href="/wiki/Mutation#Distribution_of_fitness_effects_(DFE)" title="Mutation">distribution of fitness effects (DFE)</a> for new mutations, only a minority of mutations are beneficial. Mutations with gross effects are typically deleterious. Studies in the fly <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> suggest that if a mutation changes a protein produced by a gene, this will probably be harmful, with about 70 percent of these mutations having damaging effects, and the remainder being either neutral or weakly beneficial.<sup id="cite_ref-SawyerParsch2007_28-0" class="reference"><a href="#cite_note-SawyerParsch2007-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>This biological process of mutation is represented in population-genetic models in one of two ways, either as a deterministic pressure of recurrent mutation on allele frequencies, or a source of variation. In deterministic theory, evolution begins with a predetermined set of alleles and proceeds by shifts in continuous frequencies, as if the population is infinite. The occurrence of mutations in individuals is represented by a population-level "force" or "pressure" of mutation, i.e., the force of innumerable events of mutation with a scaled magnitude u applied to shifting frequencies f(A1) to f(A2). For instance, in the classic <a href="/wiki/Mutation%E2%80%93selection_balance" title="Mutation–selection balance">mutation–selection balance</a> model,<sup id="cite_ref-CrowKimura1970_29-0" class="reference"><a href="#cite_note-CrowKimura1970-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> the force of mutation pressure pushes the frequency of an allele upward, and selection against its deleterious effects pushes the frequency downward, so that a balance is reached at equilibrium, given (in the simplest case) by f = u/s. </p><p>This concept of mutation pressure is mostly useful for considering the implications of deleterious mutation, such as the mutation load and its implications for the evolution of the mutation rate.<sup id="cite_ref-Lynch2010_30-0" class="reference"><a href="#cite_note-Lynch2010-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Transformation of populations by mutation pressure is unlikely. Haldane<sup id="cite_ref-Haldane1932_31-0" class="reference"><a href="#cite_note-Haldane1932-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> argued that it would require high mutation rates unopposed by selection, and Kimura<sup id="cite_ref-Kimura1980_32-0" class="reference"><a href="#cite_note-Kimura1980-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> concluded even more pessimistically that even this was unlikely, as the process would take too long (see <a href="/wiki/Bias_in_the_introduction_of_variation#Evolution_by_mutation_pressure" title="Bias in the introduction of variation"> evolution by mutation pressure</a>). </p><p>However, <a href="/wiki/Bias_in_the_introduction_of_variation#Evolution_by_mutation_pressure" title="Bias in the introduction of variation"> evolution by mutation pressure</a> is possible under some circumstances and has long been suggested as a possible cause for the loss of unused traits.<sup id="cite_ref-Haldane1933_33-0" class="reference"><a href="#cite_note-Haldane1933-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/Pigment" title="Pigment">pigments</a> are no longer useful when animals live in the darkness of caves, and tend to be lost.<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> An experimental example involves the loss of sporulation in experimental populations of <i>B. subtilis</i>. Sporulation is a complex trait encoded by many loci, such that the mutation rate for loss of the trait was estimated as an unusually high value, <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \mu =0.003}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>μ</mi> <mo>=</mo> <mn>0.003</mn> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \mu =0.003}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/c666ca48d0fd2c34ac0a0c70659c878861c3511f" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:9.797ex; height:2.676ex;" alt="{\displaystyle \mu =0.003}"></span>.<sup id="cite_ref-Maughan2007_35-0" class="reference"><a href="#cite_note-Maughan2007-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Loss of sporulation in this case can occur by recurrent mutation, without requiring selection for the loss of sporulation ability. When there is no selection for loss of function, the speed at which loss evolves depends more on the mutation rate than it does on the <a href="/wiki/Effective_population_size" title="Effective population size">effective population size</a>,<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> indicating that it is driven more by mutation than by genetic drift. </p><p>The role of mutation as a source of novelty is different from these classical models of mutation pressure. When population-genetic models include a rate-dependent process of mutational introduction or origination, i.e., a process that introduces new alleles including neutral and beneficial ones, then the properties of mutation may have a more direct impact on the rate and direction of evolution, even if the rate of mutation is very low.<sup id="cite_ref-Gomez2020_37-0" class="reference"><a href="#cite_note-Gomez2020-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cano2022_38-0" class="reference"><a href="#cite_note-Cano2022-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> That is, the spectrum of mutation may become very important, particularly <a href="/wiki/Mutation_bias" title="Mutation bias">mutation biases</a>, predictable differences in the rates of occurrence for different types of mutations, because <a href="/wiki/Bias_in_the_introduction_of_variation" title="Bias in the introduction of variation">bias in the introduction of variation</a> can impose biases on the course of evolution.<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> </p><p>Mutation plays a key role in other classical and recent theories including <a href="/wiki/Muller%27s_ratchet" title="Muller's ratchet">Muller's ratchet</a>, <a href="/wiki/Subfunctionalization" title="Subfunctionalization">subfunctionalization</a>, Eigen's concept of an <a href="/wiki/Error_catastrophe" title="Error catastrophe">error catastrophe</a> and Lynch's <a href="/wiki/Evolution_of_biological_complexity#Mutational_hazard_hypothesis" title="Evolution of biological complexity">mutational hazard hypothesis</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_drift">Genetic drift</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=9" title="Edit section: Genetic drift"><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/Genetic_drift" title="Genetic drift">Genetic drift</a></div> <p><i>Genetic drift</i> is a change in <a href="/wiki/Allele_frequency" title="Allele frequency">allele frequencies</a> caused by <a href="/wiki/Sampling_(statistics)" title="Sampling (statistics)">random sampling</a>.<sup id="cite_ref-Masel_2011_40-0" class="reference"><a href="#cite_note-Masel_2011-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> That is, the alleles in the offspring are a random sample of those in the parents.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Genetic drift may cause gene variants to disappear completely, and thereby reduce genetic variability. In contrast to natural selection, which makes gene variants more common or less common depending on their reproductive success,<sup id="cite_ref-avers_42-0" class="reference"><a href="#cite_note-avers-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> the changes due to genetic drift are not driven by environmental or adaptive pressures, and are equally likely to make an allele more common as less common. </p><p>The effect of genetic drift is larger for alleles present in few copies than when an allele is present in many copies. The population genetics of genetic drift are described using either <a href="/wiki/Branching_process" title="Branching process">branching processes</a> or a <a href="/wiki/Diffusion_equation" title="Diffusion equation">diffusion equation</a> describing changes in allele frequency.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> These approaches are usually applied to the Wright-Fisher and <a href="/wiki/Moran_process" title="Moran process">Moran</a> models of population genetics. Assuming genetic drift is the only evolutionary force acting on an allele, after t generations in many replicated populations, starting with allele frequencies of p and q, the variance in allele frequency across those populations is </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle V_{t}\approx pq\left(1-\exp \left\{-{\frac {t}{2N_{e}}}\right\}\right).}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>V</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>t</mi> </mrow> </msub> <mo>≈</mo> <mi>p</mi> <mi>q</mi> <mrow> <mo>(</mo> <mrow> <mn>1</mn> <mo>−</mo> <mi>exp</mi> <mo>⁡</mo> <mrow> <mo>{</mo> <mrow> <mo>−</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mi>t</mi> <mrow> <mn>2</mn> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>e</mi> </mrow> </msub> </mrow> </mfrac> </mrow> </mrow> <mo>}</mo> </mrow> </mrow> <mo>)</mo> </mrow> <mo>.</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle V_{t}\approx pq\left(1-\exp \left\{-{\frac {t}{2N_{e}}}\right\}\right).}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/3dfd633cb35bee9cf9364b02a12437a0d0e6c55e" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.505ex; width:30.074ex; height:6.176ex;" alt="{\displaystyle V_{t}\approx pq\left(1-\exp \left\{-{\frac {t}{2N_{e}}}\right\}\right).}"></span><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup></dd></dl> <p><a href="/wiki/Ronald_Fisher" title="Ronald Fisher">Ronald Fisher</a> held the view that genetic drift plays at the most a minor role in evolution, and this remained the dominant view for several decades. No population genetics perspective have ever given genetic drift a central role by itself, but some have made genetic drift important in combination with another non-selective force. The <a href="/wiki/Shifting_balance_theory" title="Shifting balance theory">shifting balance theory</a> of <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a> held that the combination of population structure and genetic drift was important. <a href="/wiki/Motoo_Kimura" title="Motoo Kimura">Motoo Kimura</a>'s <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of molecular evolution</a> claims that most genetic differences within and between populations are caused by the combination of neutral mutations and genetic drift.<sup id="cite_ref-Futuyma_1998_320_45-0" class="reference"><a href="#cite_note-Futuyma_1998_320-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>The role of genetic drift by means of <a href="/wiki/Sampling_error" title="Sampling error">sampling error</a> in evolution has been criticized by <a href="/wiki/John_H_Gillespie" class="mw-redirect" title="John H Gillespie">John H Gillespie</a><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Will_Provine" class="mw-redirect" title="Will Provine">Will Provine</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> who argue that selection on linked sites is a more important stochastic force, doing the work traditionally ascribed to genetic drift by means of sampling error. The mathematical properties of genetic draft are different from those of genetic drift.<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> The direction of the random change in allele frequency is <a href="/wiki/Autocorrelation" title="Autocorrelation">autocorrelated</a> across generations.<sup id="cite_ref-Masel_2011_40-1" class="reference"><a href="#cite_note-Masel_2011-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Gene_flow">Gene flow</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=10" title="Edit section: Gene flow"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Gene_flow_final.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Gene_flow_final.png/220px-Gene_flow_final.png" decoding="async" width="220" height="104" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Gene_flow_final.png/330px-Gene_flow_final.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Gene_flow_final.png/440px-Gene_flow_final.png 2x" data-file-width="1531" data-file-height="724" /></a><figcaption>Gene flow is the transfer of <a href="/wiki/Allele" title="Allele">alleles</a> from one <a href="/wiki/Population" title="Population">population</a> to another population through immigration of individuals. In this example, one of the birds from population A <a href="/wiki/Immigrate" class="mw-redirect" title="Immigrate">immigrates</a> to population B, which has fewer of the dominant alleles, and through mating incorporates its alleles into the other population.</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/Gene_flow" title="Gene flow">Gene flow</a></div> <p>Because of physical barriers to migration, along with the limited tendency for individuals to move or spread (<a href="https://en.wiktionary.org/wiki/vagility" class="extiw" title="wikt:vagility">vagility</a>), and tendency to remain or come back to natal place (<a href="/wiki/Philopatry" title="Philopatry">philopatry</a>), natural populations rarely all interbreed as may be assumed in theoretical random models (<a href="/wiki/Panmixy" class="mw-redirect" title="Panmixy">panmixy</a>).<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> There is usually a geographic range within which individuals are more closely <a href="/wiki/Coefficient_of_relatedness" class="mw-redirect" title="Coefficient of relatedness">related</a> to one another than those randomly selected from the general population. This is described as the extent to which a population is genetically structured.<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> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Greatwall_large.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Greatwall_large.jpg/220px-Greatwall_large.jpg" decoding="async" width="220" height="254" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Greatwall_large.jpg/330px-Greatwall_large.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Greatwall_large.jpg/440px-Greatwall_large.jpg 2x" data-file-width="1275" data-file-height="1473" /></a><figcaption>The <a href="/wiki/Great_Wall_of_China" title="Great Wall of China">Great Wall of China</a> is an obstacle to gene flow of some terrestrial species.<sup id="cite_ref-Su_Qu_2003_51-0" class="reference"><a href="#cite_note-Su_Qu_2003-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Genetic structuring can be caused by migration due to historical <a href="/wiki/Climate_change_(general_concept)" class="mw-redirect" title="Climate change (general concept)">climate change</a>, species <a href="/wiki/Range_expansion" class="mw-redirect" title="Range expansion">range expansion</a> or current availability of <a href="/wiki/Habitat" title="Habitat">habitat</a>. Gene flow is hindered by mountain ranges, oceans and deserts or even human-made structures such as the <a href="/wiki/Great_Wall_of_China" title="Great Wall of China">Great Wall of China</a>, which has hindered the flow of plant genes.<sup id="cite_ref-Su_Qu_2003_51-1" class="reference"><a href="#cite_note-Su_Qu_2003-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a> is the exchange of genes between populations or species, breaking down the structure. Examples of gene flow within a species include the migration and then breeding of organisms, or the exchange of <a href="/wiki/Pollen" title="Pollen">pollen</a>. Gene transfer between species includes the formation of <a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">hybrid</a> organisms and <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>. Population genetic models can be used to identify which populations show significant genetic isolation from one another, and to reconstruct their history.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>Subjecting a population to isolation leads to <a href="/wiki/Inbreeding_depression" title="Inbreeding depression">inbreeding depression</a>. Migration into a population can introduce new genetic variants,<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> potentially contributing to <a href="/wiki/Evolutionary_rescue" title="Evolutionary rescue">evolutionary rescue</a>. If a significant proportion of individuals or gametes migrate, it can also change allele frequencies, e.g. giving rise to <a href="/wiki/Genetic_load#Migration_load" title="Genetic load">migration load</a>.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>In the presence of gene flow, other <a href="/wiki/Reproductive_isolation" title="Reproductive isolation">barriers to hybridization</a> between two diverging populations of an <a href="/wiki/Outcrossing" title="Outcrossing">outcrossing</a> species are required for the populations to <a href="/wiki/Speciation" title="Speciation">become new species</a>. </p> <div class="mw-heading mw-heading4"><h4 id="Horizontal_gene_transfer">Horizontal gene transfer</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=11" title="Edit section: Horizontal gene transfer"><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:Tree_Of_Life_(with_horizontal_gene_transfer).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/220px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/330px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/440px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 2x" data-file-width="1782" data-file-height="2370" /></a><figcaption>Current tree of life showing vertical and horizontal gene transfers</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/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></div> <p>Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>.<sup id="cite_ref-Boucher_Douady_2003_55-0" class="reference"><a href="#cite_note-Boucher_Douady_2003-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> In medicine, this contributes to the spread of <a href="/wiki/Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a>, as when one bacteria acquires resistance genes it can rapidly transfer them to other species.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> Horizontal transfer of genes from bacteria to eukaryotes such as the yeast <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i> and the adzuki bean beetle <i><a href="/wiki/Callosobruchus_chinensis" title="Callosobruchus chinensis">Callosobruchus chinensis</a></i> may also have occurred.<sup id="cite_ref-Kondo_Nikoh_2002_57-0" class="reference"><a href="#cite_note-Kondo_Nikoh_2002-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup><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> An example of larger-scale transfers are the eukaryotic <a href="/wiki/Bdelloidea" title="Bdelloidea">bdelloid rotifers</a>, which appear to have received a range of genes from bacteria, fungi, and plants.<sup id="cite_ref-Gladyshev_Meselson_2008_59-0" class="reference"><a href="#cite_note-Gladyshev_Meselson_2008-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Virus" title="Virus">Viruses</a> can also carry DNA between organisms, allowing transfer of genes even across <a href="/wiki/Domain_(biology)" title="Domain (biology)">biological domains</a>.<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> Large-scale gene transfer has also occurred between the ancestors of <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic cells</a> and prokaryotes, during the acquisition of <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a> and <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a>.<sup id="cite_ref-rgruqh_61-0" class="reference"><a href="#cite_note-rgruqh-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Linkage">Linkage</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=12" title="Edit section: Linkage"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>If all genes are in <a href="/wiki/Linkage_equilibrium" class="mw-redirect" title="Linkage equilibrium">linkage equilibrium</a>, the effect of an allele at one locus can be averaged across the <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a> at other loci. In reality, one allele is frequently found in <a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a> with genes at other loci, especially with genes located nearby on the same chromosome. <a href="/wiki/Recombination_(biology)" class="mw-redirect" title="Recombination (biology)">Recombination</a> breaks up this linkage disequilibrium too slowly to avoid <a href="/wiki/Genetic_hitchhiking" title="Genetic hitchhiking">genetic hitchhiking</a>, where an allele at one locus rises to high frequency because it is <a href="/wiki/Genetic_linkage" title="Genetic linkage">linked</a> to an allele under selection at a nearby locus. Linkage also slows down the rate of adaptation, even in sexual populations.<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><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 effect of linkage disequilibrium in slowing down the rate of adaptive evolution arises from a combination of the <a href="/wiki/Hill%E2%80%93Robertson_effect" title="Hill–Robertson effect">Hill–Robertson effect</a> (delays in bringing beneficial mutations together) and <a href="/wiki/Background_selection" title="Background selection">background selection</a> (delays in separating beneficial mutations from deleterious <a href="/wiki/Genetic_hitchhiking" title="Genetic hitchhiking">hitchhikers</a>). </p><p>Linkage is a problem for population genetic models that treat one gene locus at a time. It can, however, be exploited as a method for detecting the action of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> via <a href="/wiki/Selective_sweep" title="Selective sweep">selective sweeps</a>. </p><p>In the extreme case of an <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexual population</a>, linkage is complete, and population genetic equations can be derived and solved in terms of a travelling <a href="/wiki/Wave" title="Wave">wave</a> of genotype frequencies along a simple <a href="/wiki/Fitness_landscape" title="Fitness landscape">fitness landscape</a>.<sup id="cite_ref-Desai_65-0" class="reference"><a href="#cite_note-Desai-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Most <a href="/wiki/Microorganisms" class="mw-redirect" title="Microorganisms">microbes</a>, such as <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, are asexual. The population genetics of their <a href="/wiki/Adaptation" title="Adaptation">adaptation</a> have two contrasting regimes. When the product of the beneficial mutation rate and population size is small, asexual populations follow a "successional regime" of origin-fixation dynamics, with adaptation rate strongly dependent on this product. When the product is much larger, asexual populations follow a "concurrent mutations" regime with adaptation rate less dependent on the product, characterized by <a href="/wiki/Clonal_interference" title="Clonal interference">clonal interference</a> and the appearance of a new beneficial mutation before the last one has <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixed</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=13" title="Edit section: Applications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Explaining_levels_of_genetic_variation">Explaining levels of genetic variation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=14" title="Edit section: Explaining levels of genetic variation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">Neutral theory</a> predicts that the level of <a href="/wiki/Nucleotide_diversity" title="Nucleotide diversity">nucleotide diversity</a> in a population will be proportional to the product of the population size and the neutral mutation rate. The fact that levels of genetic diversity vary much less than population sizes do is known as the "paradox of variation".<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> While high levels of genetic diversity were one of the original arguments in favor of neutral theory, the paradox of variation has been one of the strongest arguments against neutral theory. </p><p>It is clear that levels of genetic diversity vary greatly within a species as a function of local recombination rate, due to both <a href="/wiki/Genetic_hitchhiking" title="Genetic hitchhiking">genetic hitchhiking</a> and <a href="/wiki/Background_selection" title="Background selection">background selection</a>. Most current solutions to the paradox of variation invoke some level of selection at linked sites.<sup id="cite_ref-ellegren_67-0" class="reference"><a href="#cite_note-ellegren-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> For example, one analysis suggests that larger populations have more selective sweeps, which remove more neutral genetic diversity.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> A negative correlation between mutation rate and population size may also contribute.<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> </p><p>Life history affects genetic diversity more than population history does, e.g. <a href="/wiki/R/K_selection_theory" title="R/K selection theory">r-strategists</a> have more genetic diversity.<sup id="cite_ref-ellegren_67-1" class="reference"><a href="#cite_note-ellegren-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Detecting_selection">Detecting selection</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=15" title="Edit section: Detecting selection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Population genetics models are used to infer which genes are undergoing selection. One common approach is to look for regions of high <a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a> and low genetic variance along the chromosome, to detect recent <a href="/wiki/Selective_sweep" title="Selective sweep">selective sweeps</a>. </p><p>A second common approach is the <a href="/wiki/McDonald%E2%80%93Kreitman_test" title="McDonald–Kreitman test">McDonald–Kreitman test</a> which compares the amount of variation within a species (<a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">polymorphism</a>) to the divergence between species (substitutions) at two types of sites; one assumed to be neutral. Typically, <a href="/wiki/Synonymous_substitution" title="Synonymous substitution">synonymous</a> sites are assumed to be neutral.<sup id="cite_ref-charlesworth_70-0" class="reference"><a href="#cite_note-charlesworth-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> Genes undergoing positive selection have an excess of divergent sites relative to polymorphic sites. The test can also be used to obtain a genome-wide estimate of the proportion of substitutions that are fixed by positive selection, α.<sup id="cite_ref-eyrewalker_71-0" class="reference"><a href="#cite_note-eyrewalker-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-alpha_72-0" class="reference"><a href="#cite_note-alpha-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> According to the <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of molecular evolution</a>, this number should be near zero. High numbers have therefore been interpreted as a genome-wide falsification of neutral theory.<sup id="cite_ref-Hahn_73-0" class="reference"><a href="#cite_note-Hahn-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Demographic_inference">Demographic inference</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=16" title="Edit section: Demographic inference"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The simplest test for population structure in a sexually reproducing, diploid species, is to see whether genotype frequencies follow Hardy-Weinberg proportions as a function of allele frequencies. For example, in the simplest case of a single locus with two <a href="/wiki/Allele" title="Allele">alleles</a> denoted <b>A</b> and <b>a</b> at frequencies <i>p</i> and <i>q</i>, random mating predicts freq(<b>AA</b>) = <i>p</i><sup>2</sup> for the <b>AA</b> <a href="/wiki/Homozygote" class="mw-redirect" title="Homozygote">homozygotes</a>, freq(<b>aa</b>) = <i>q</i><sup>2</sup> for the <b>aa</b> homozygotes, and freq(<b>Aa</b>) = 2<i>pq</i> for the <a href="/wiki/Heterozygote" class="mw-redirect" title="Heterozygote">heterozygotes</a>. In the absence of population structure, Hardy-Weinberg proportions are reached within 1–2 generations of random mating. More typically, there is an excess of homozygotes, indicative of population structure. The extent of this excess can be quantified as the <a href="/wiki/F-statistics" title="F-statistics">inbreeding coefficient, F</a>. </p><p>Individuals can be clustered into <i>K</i> subpopulations.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup><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> The degree of population structure can then be calculated using <a href="/wiki/Fixation_index" title="Fixation index">F<sub>ST</sub></a>, which is a measure of the proportion of genetic variance that can be explained by population structure. Genetic population structure can then be related to geographic structure, and <a href="/wiki/Genetic_admixture" title="Genetic admixture">genetic admixture</a> can be detected. </p><p><a href="/wiki/Coalescent_theory" title="Coalescent theory">Coalescent theory</a> relates genetic diversity in a sample to demographic history of the population from which it was taken. It normally assumes <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutrality</a>, and so sequences from more neutrally evolving portions of genomes are therefore selected for such analyses. It can be used to infer the relationships between species (<a href="/wiki/Computational_phylogenetics" title="Computational phylogenetics">phylogenetics</a>), as well as the population structure, demographic history (e.g. <a href="/wiki/Population_bottlenecks" class="mw-redirect" title="Population bottlenecks">population bottlenecks</a>, <a href="/wiki/Population_growth" title="Population growth">population growth</a>), <a href="/wiki/Biological_dispersal" title="Biological dispersal">biological dispersal</a>, <a href="/wiki/Source%E2%80%93sink_dynamics" title="Source–sink dynamics">source–sink dynamics</a><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> and <a href="/wiki/Introgression" title="Introgression">introgression</a> within a species. </p><p>Another approach to demographic inference relies on the <a href="/wiki/Allele_frequency_spectrum" title="Allele frequency spectrum">allele frequency spectrum</a>.<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="Evolution_of_genetic_systems">Evolution of genetic systems</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Population_genetics&action=edit&section=17" title="Edit section: Evolution of genetic systems"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By assuming that there are loci that control the genetic system itself, population genetic models are created to describe the <a href="/wiki/Evolution_of_dominance" class="mw-redirect" title="Evolution of dominance">evolution of dominance</a> and other forms of <a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">robustness</a>, the <a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">evolution of sexual reproduction</a> and recombination rates, the evolution of <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rates</a>, the evolution of <a href="/wiki/Evolutionary_capacitance" title="Evolutionary capacitance">evolutionary capacitors</a>, the evolution of <a href="/wiki/Signalling_theory" title="Signalling theory">costly signalling traits</a>, the <a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">evolution of ageing</a>, and the evolution of <a href="/wiki/Co-operation_(evolution)" class="mw-redirect" title="Co-operation (evolution)">co-operation</a>. For example, most mutations are deleterious, so the optimal <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> for a species may be a trade-off between the damage from a high deleterious mutation rate and the <a href="/wiki/Metabolism" title="Metabolism">metabolic</a> costs of maintaining systems to reduce the mutation rate, such as DNA repair enzymes.<sup id="cite_ref-Sniegowski_78-0" class="reference"><a href="#cite_note-Sniegowski-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p><p>One important aspect of such models is that selection is only strong enough to purge deleterious mutations and hence overpower mutational bias towards degradation if the selection coefficient s is greater than the inverse of the <a href="/wiki/Effective_population_size" title="Effective population size">effective population size</a>. This is known as the drift barrier and is related to the <a href="/wiki/Nearly_neutral_theory_of_molecular_evolution" title="Nearly neutral theory of molecular evolution">nearly neutral theory of molecular evolution</a>. Drift barrier theory predicts that species with large effective population sizes will have highly streamlined, efficient genetic systems, while those with small population sizes will have bloated and complex <a href="/wiki/Genome" title="Genome">genomes</a> containing for example <a href="/wiki/Intron" title="Intron">introns</a> and <a href="/wiki/Transposable_element" title="Transposable element">transposable elements</a>.<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> However, somewhat paradoxically, species with large population sizes might be so tolerant to the consequences of certain types of errors that they evolve higher error rates, e.g. in <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> and <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, than small populations.<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> </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=Population_genetics&action=edit&section=18" 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: 30em;"> <ul><li><a href="/wiki/Allele_frequency_spectrum" title="Allele frequency spectrum">Allele frequency spectrum</a></li> <li><a href="/wiki/Assortative_mating" title="Assortative mating">Assortative mating</a></li> <li><a href="/wiki/Balancing_selection" title="Balancing selection">Balancing selection</a></li> <li><a href="/wiki/Bias_in_the_introduction_of_variation" title="Bias in the introduction of variation">Bias in the introduction of variation</a></li> <li><a href="/wiki/Coalescent_theory" title="Coalescent theory">Coalescent theory</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation genetics</a></li> <li><a href="/wiki/Constructive_neutral_evolution" title="Constructive neutral evolution">Constructive neutral evolution</a></li> <li><a href="/wiki/Directional_selection" title="Directional selection">Directional selection</a></li> <li><a href="/wiki/Mutation#Distribution_of_fitness_effects_(DFE)" title="Mutation">Distribution of fitness effects (DFE)</a></li> <li><a href="/wiki/Drift-barrier_hypothesis" title="Drift-barrier hypothesis">Drift-barrier hypothesis</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Error_catastrophe" title="Error catastrophe">Error catastrophe</a></li> <li><a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">Evolutionary landscape</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a></li> <li><a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">Fixation</a></li> <li><a href="/wiki/Founder_effect" title="Founder effect">Founder effect</a></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Gene_pool" title="Gene pool">Gene pool</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Genetic_hitchhiking" title="Genetic hitchhiking">Genetic hitchhiking</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Haldane%27s_dilemma" title="Haldane's dilemma">Haldane's dilemma</a></li> <li><a href="/wiki/Hardy%E2%80%93Weinberg_principle" title="Hardy–Weinberg principle">Hardy-Weinberg equilibrium</a></li> <li><a href="/wiki/Idealised_population" title="Idealised population">Idealised population</a></li> <li><a href="/wiki/Inbreeding" title="Inbreeding">Inbreeding</a></li> <li><a href="/wiki/Inbreeding_depression" title="Inbreeding depression">Inbreeding depression</a></li> <li><a href="/wiki/Infinite_sites_model" title="Infinite sites model">Infinite sites model</a></li> <li><a href="/wiki/Intragenomic_conflict" title="Intragenomic conflict">Intragenomic conflict</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">Linkage disequilibrium</a></li> <li><a href="/wiki/McDonald%E2%80%93Kreitman_test" title="McDonald–Kreitman test">McDonald–Kreitman test</a></li> <li><a href="/wiki/Meiotic_drive" title="Meiotic drive">Meiotic drive</a></li> <li><a href="/wiki/Muller%27s_ratchet" title="Muller's ratchet">Muller's ratchet</a></li> <li><a href="/wiki/Mutation%E2%80%93selection_balance" title="Mutation–selection balance">Mutation–selection balance</a></li> <li><a href="/wiki/Evolution_of_biological_complexity#Mutational_hazard_hypothesis" title="Evolution of biological complexity">Mutational hazard hypothesis</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">Neutral theory of molecular evolution</a></li> <li><a href="/wiki/Panmixia" title="Panmixia">Panmixia</a></li> <li><a href="/wiki/Outbreeding_depression" title="Outbreeding depression">Outbreeding depression</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Polygenic_adaptation" title="Polygenic adaptation">Polygenic adaptation</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Fisherian_runaway" title="Fisherian runaway">Runaway process (Fisherian runaway)</a></li> <li><a href="/wiki/Selective_sweep" title="Selective sweep">Selective sweep</a></li> <li><a href="/wiki/Selfish_genetic_element" title="Selfish genetic element">Selfish genetic element</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a></li> <li><a href="/wiki/Shifting_balance_theory" title="Shifting balance theory">Shifting balance theory</a></li> <li><a href="/wiki/Subfunctionalization" title="Subfunctionalization">Subfunctionalization</a></li> <li><a href="/wiki/Supergene" title="Supergene">Supergene (co-adapted gene complex)</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=Population_genetics&action=edit&section=19" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></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 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.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 class="citation web cs1"><a rel="nofollow" class="external text" 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Stearns">Stephen C. Stearns</a> (<a href="/wiki/Yale_University" title="Yale University">Yale University</a>)</li> <li><a href="/wiki/National_Geographic_Society" title="National Geographic Society">National Geographic</a>: <a rel="nofollow" class="external text" href="https://web.archive.org/web/20060326213734/https://www9.nationalgeographic.com/genographic/atlas.html">Atlas of the Human Journey</a> (<a href="/wiki/Haplogroup" title="Haplogroup">Haplogroup</a>-based human migration maps)</li></ul> <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 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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:Population_genetics" title="Template:Population genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Population_genetics" title="Template talk:Population genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Population_genetics" title="Special:EditPage/Template:Population genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Population_genetics" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Population genetics</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Key concepts</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/Hardy%E2%80%93Weinberg_principle" title="Hardy–Weinberg principle">Hardy–Weinberg principle</a></li> <li><a href="/wiki/Genetic_linkage" title="Genetic linkage">Genetic linkage</a></li> <li><a href="/wiki/Identity_by_descent" title="Identity by descent">Identity by descent</a></li> <li><a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">Linkage disequilibrium</a></li> <li><a href="/wiki/Fisher%27s_fundamental_theorem_of_natural_selection" title="Fisher's fundamental theorem of natural selection">Fisher's fundamental theorem</a></li> <li><a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">Neutral theory</a></li> <li><a href="/wiki/Shifting_balance_theory" title="Shifting balance theory">Shifting balance theory</a></li> <li><a href="/wiki/Price_equation" title="Price equation">Price equation</a></li> <li><a href="/wiki/Coefficient_of_inbreeding" title="Coefficient of inbreeding">Coefficient of inbreeding</a></li> <li><a href="/wiki/Coefficient_of_relationship" title="Coefficient of relationship">Coefficient of relationship</a></li> <li><a href="/wiki/Selection_coefficient" title="Selection coefficient">Selection coefficient</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Population_structure_(genetics)" title="Population structure (genetics)">Population structure</a></li> <li><a href="/wiki/Constructive_neutral_evolution" title="Constructive neutral evolution">Constructive neutral evolution</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/Natural_selection" title="Natural selection">Selection</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Natural_selection" title="Natural selection">Natural</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual</a></li> <li><a href="/wiki/Ecological_selection" class="mw-redirect" title="Ecological selection">Ecological</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><div style="display: inline-block; line-height: 1.2em; padding: .1em 0; line-height:1.1em;">Effects of selection<br />on genomic variation</div></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_hitchhiking" title="Genetic hitchhiking">Genetic hitchhiking</a></li> <li><a href="/wiki/Negative_selection_(natural_selection)" title="Negative selection (natural selection)">Background selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Population_bottleneck" title="Population bottleneck">Population bottleneck</a></li> <li><a href="/wiki/Founder_effect" title="Founder effect">Founder effect</a></li> <li><a href="/wiki/Coalescent_theory" title="Coalescent theory">Coalescence</a></li> <li><a href="/wiki/Balding%E2%80%93Nichols_model" title="Balding–Nichols model">Balding–Nichols model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Founders</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/Ronald_Fisher" title="Ronald Fisher">R. A. Fisher</a></li> <li><a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a></li> <li><a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Evolution" title="Evolution">Evolution</a></li> <li><a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Landscape_genetics" title="Landscape genetics">Landscape genetics</a> and <a href="/wiki/Landscape_genomics" title="Landscape genomics">genomics</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Phylogeography" title="Phylogeography">Phylogeography</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align:center;"><div> <ul><li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</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="Human_genetics" 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:Human_genetics" title="Template:Human genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Human_genetics" title="Template talk:Human genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Human_genetics" title="Special:EditPage/Template:Human genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_genetics" title="Human genetics">Human genetics</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Sub-topics</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/Human_genome" title="Human genome">Human genome</a> <ul><li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li></ul></li> <li><a href="/wiki/Human_evolutionary_genetics" title="Human evolutionary genetics">Evolutionary genetics</a> <ul><li><a href="/wiki/Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Human-chimp MRCA</a></li> <li><a href="/wiki/Neanderthal_genetics" title="Neanderthal genetics">Neanderthal genetics</a> <ul><li><a href="/wiki/Neanderthal_genome_project" title="Neanderthal genome project">Neanderthal genome project</a></li></ul></li> <li><a href="/wiki/Timeline_of_human_evolution" title="Timeline of human evolution">Timeline</a></li></ul></li> <li><a href="/wiki/Human_genetic_variation" title="Human genetic variation">Genetic variation</a> <ul><li><a href="/wiki/Blood_type_distribution_by_country" title="Blood type distribution by country">Blood type distribution by country</a></li> <li><a href="/wiki/Genealogical_DNA_test" title="Genealogical DNA test">Genealogical DNA test</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Race_and_genetics" title="Race and genetics">Race and genetics</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent evolution</a></li> <li><a href="/wiki/Surname_DNA_project" title="Surname DNA project">Surname DNA project</a></li></ul></li> <li><a href="/wiki/Human_genetic_enhancement" title="Human genetic enhancement">Genetic enhancement</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Genetic history</a> <br />by region</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/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a> <ul><li><a href="/wiki/Genetic_history_of_North_Africa" title="Genetic history of North Africa">North Africa</a></li> <li><a href="/wiki/Genetic_history_of_West_Africa" title="Genetic history of West Africa">West Africa‎‎</a></li> <li><a href="/wiki/Genetic_history_of_Central_Africa" title="Genetic history of Central Africa">Central Africa</a></li> <li><a href="/wiki/Genetic_history_of_Eastern_Africa" title="Genetic history of Eastern Africa">Eastern Africa‎‎</a></li> <li><a href="/wiki/Genetic_history_of_Southern_Africa" title="Genetic history of Southern Africa">Southern Africa</a></li> <li><a href="/wiki/Genetic_history_of_the_African_diaspora" title="Genetic history of the African diaspora">African diaspora</a></li></ul></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a> <ul><li><a href="/wiki/Peopling_of_India" title="Peopling of India">India</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">Middle East</a> <ul><li><a href="/wiki/Early_Anatolian_farmers" class="mw-redirect" title="Early Anatolian farmers">Early Anatolian farmers</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Caucasus" class="mw-redirect" title="Genetic history of the Caucasus">Caucasus</a> <ul><li><a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Caucasus hunter-gatherer</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a> <ul><li><a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmers</a></li> <li><a href="/wiki/Western_hunter-gatherer" title="Western hunter-gatherer">Western hunter-gatherer</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">British Isles</a></li> <li><a href="/wiki/Genetic_history_of_the_Iberian_Peninsula" title="Genetic history of the Iberian Peninsula">Iberia</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Eastern_hunter-gatherer" title="Eastern hunter-gatherer">Eastern hunter-gatherer</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Central_Asia" class="mw-redirect" title="Genetic history of Central Asia">Central Asia</a> <ul><li><a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_East_Asians" title="Genetic history of East Asians">East Asia</a> <ul><li><a href="/wiki/Ancient_Northeast_Asian" title="Ancient Northeast Asian">Ancient Northeast Asian</a></li> <li><a href="/wiki/Ancient_Paleo-Siberian" title="Ancient Paleo-Siberian">Ancient Paleo-Siberian</a></li> <li><a href="/wiki/Peopling_of_China" title="Peopling of China">China</a></li></ul></li> <li><a href="/wiki/Peopling_of_Southeast_Asia" title="Peopling of Southeast Asia">Southeast Asia</a> <ul><li><a href="/wiki/Peopling_of_Thailand" title="Peopling of Thailand">Thailand</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of Indigenous peoples of the Americas">America</a> <ul><li><a href="/wiki/Ancient_Beringian" title="Ancient Beringian">Ancient Beringian</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a class="mw-selflink selflink">Population genetics</a> <br />by group</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Europe <ul><li><a href="/wiki/Origin_of_the_Basques" title="Origin of the Basques">Basques</a></li> <li><a href="/wiki/Genetic_studies_on_Bosniaks" title="Genetic studies on Bosniaks">Bosniaks</a></li> <li><a href="/wiki/Genetic_studies_on_Bulgarians" title="Genetic studies on Bulgarians">Bulgarians</a></li> <li><a href="/wiki/Genetic_studies_on_Croats" title="Genetic studies on Croats">Croats</a></li> <li><a href="/wiki/Genetic_studies_on_Romanians" class="mw-redirect" title="Genetic studies on Romanians">Romanians</a></li> <li><a href="/wiki/Genetic_studies_on_Russians" title="Genetic studies on Russians">Russians</a></li> <li><a href="/wiki/Genetic_studies_on_Sami" title="Genetic studies on Sami">Sami</a></li> <li><a href="/wiki/Genetic_studies_on_Serbs" title="Genetic studies on Serbs">Serbs</a></li></ul></li> <li>MENA <ul><li><a href="/wiki/Genetic_studies_on_Arabs" title="Genetic studies on Arabs">Arabs</a></li> <li><a href="/wiki/Origin_of_the_Azerbaijanis" title="Origin of the Azerbaijanis">Azerbaijanis</a></li> <li><a href="/wiki/Genetic_history_of_Egypt" title="Genetic history of Egypt">Egyptians</a></li> <li><a href="/wiki/Genetic_studies_on_Jews" class="mw-redirect" title="Genetic studies on Jews">Jews</a></li> <li><a href="/wiki/Genetic_studies_on_Moroccans" title="Genetic studies on Moroccans">Moroccans</a></li> <li><a href="/wiki/Genetic_studies_on_Turkish_people" title="Genetic studies on Turkish people">Turks</a></li></ul></li> <li>South Asia <ul><li><a href="/wiki/Genetic_studies_on_Gujarati_people" title="Genetic studies on Gujarati people">Gujaratis</a></li> <li><a href="/wiki/Genetic_studies_on_Sinhalese" title="Genetic studies on Sinhalese">Sinhalese</a></li> <li><a href="/wiki/Genetic_studies_on_Sri_Lankan_Tamils" title="Genetic studies on Sri Lankan Tamils">Tamils (Sri Lankan)</a></li></ul></li> <li>East Asia <ul><li><a href="/wiki/Genetic_studies_on_Han_Chinese" class="mw-redirect" title="Genetic studies on Han Chinese">Han Chinese</a></li> <li><a href="/wiki/Genetic_and_anthropometric_studies_on_Japanese_people" title="Genetic and anthropometric studies on Japanese people">Japanese</a></li></ul></li> <li>Southeast Asia <ul><li><a href="/wiki/Genetic_studies_on_Filipinos" title="Genetic studies on Filipinos">Filipinos</a></li></ul></li> <li>Sub-Saharan Africa <ul><li><a href="/wiki/Genetic_studies_on_Hutu_and_Tutsi" class="mw-redirect" title="Genetic studies on Hutu and Tutsi">Hutu/Tutsi</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><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><b><a href="/wiki/Category:Human_genetics" title="Category:Human genetics">Category</a></b></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="Genetics" 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:Genetics" title="Template:Genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetics" title="Template talk:Genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetics" title="Special:EditPage/Template:Genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Genetics" title="Genetics">Genetics</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Timeline_of_the_history_of_genetics" title="Timeline of the history of genetics">Timeline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li> <li><a href="/wiki/Glossary_of_genetics" class="mw-redirect" title="Glossary of genetics">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key components</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chromosome" 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></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a class="mw-selflink selflink">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a> of</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a></li> <li><a href="/wiki/Genetic_history_of_indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of indigenous peoples of the Americas">the Americas</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Genetic diversity</a></li> <li><a href="/wiki/Genetic_monitoring" title="Genetic monitoring">Genetic monitoring</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/He_Jiankui_affair" title="He Jiankui affair">He Jiankui affair</a></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Plant_genetics" title="Plant genetics">Plant genetics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></li> <li><a href="/wiki/List_of_genetics_research_organizations" title="List of genetics research organizations">List of genetics research organizations</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><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></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="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</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/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</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/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</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/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</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/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</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/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</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/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</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/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</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/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><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:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</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="Evolutionary_psychology" 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:Evolutionary_psychology" title="Template:Evolutionary psychology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_psychology" title="Template talk:Evolutionary psychology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_psychology" title="Special:EditPage/Template:Evolutionary psychology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_psychology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</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/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a>/<a href="/wiki/Motor_skill" title="Motor skill">skill</a></li> <li><a href="/wiki/Human_multitasking" title="Human multitasking">Multitasking</a></li> <li><a href="/wiki/Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li> <li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Eye</a></li> <li><a href="/wiki/Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</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/Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a> <ul><li><a href="/wiki/Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li> <li><a href="/wiki/Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a> <ul><li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Crime</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Language</a> <ul><li><a href="/wiki/Origin_of_language" title="Origin of language">Origin</a></li> <li><a href="/wiki/Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a> <ul><li><a href="/wiki/Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li> <li><a href="/wiki/Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a> <ul><li><a href="/wiki/Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li> <li><a href="/wiki/Cultural_universal" title="Cultural universal">Universals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Attachment_theory" title="Attachment theory">Attachment</a></li> <li><a href="/wiki/Human_bonding" title="Human bonding">Bonding</a></li> <li><a href="/wiki/Affectional_bond" title="Affectional bond">Affectional</a>/<a href="/wiki/Maternal_bond" title="Maternal bond">maternal</a>/<a href="/wiki/Paternal_bond" title="Paternal bond">paternal bond</a></li> <li><a href="/wiki/Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li> <li><a href="/wiki/Attachment_in_children" title="Attachment in children">Childhood attachment</a></li> <li><a href="/wiki/Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li> <li><a href="/wiki/Cognitive_development" title="Cognitive development">Cognitive development</a></li> <li><a href="/wiki/Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li> <li><a href="/wiki/Language_acquisition" title="Language acquisition">Language acquisition</a></li> <li><a href="/wiki/Personality_development" title="Personality development">Personality development</a></li> <li><a href="/wiki/Socialization" title="Socialization">Socialization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ergonomics" title="Ergonomics">Human factors</a> /<br /><a href="/wiki/Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</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/Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a> <ul><li><a href="/wiki/Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li> <li><a href="/wiki/Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li> <li><a href="/wiki/Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li> <li><a href="/wiki/Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li> <li><a href="/wiki/Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li> <li><a href="/wiki/Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li> <li><a href="/wiki/Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li> <li><a href="/wiki/Accident-proneness" title="Accident-proneness">Hypophobia</a></li> <li><a href="/wiki/Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li> <li><a href="/wiki/Mind-blindness" title="Mind-blindness">Mind-blindness</a></li> <li><a href="/wiki/Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li> <li><a href="/wiki/Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li> <li><a href="/wiki/Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Screen_time" title="Screen time">Screen time</a></li> <li><a href="/wiki/Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li> <li><a href="/wiki/Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li> <li><a href="/wiki/Societal_impacts_of_cars" title="Societal impacts of cars">Societal impacts of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</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/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Academic disciplines</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/Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a 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title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a 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