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class="vector-toc-numb">2</span> <span>Phenotypic variation</span> </div> </a> <button aria-controls="toc-Phenotypic_variation-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 Phenotypic variation subsection</span> </button> <ul id="toc-Phenotypic_variation-sublist" class="vector-toc-list"> <li id="toc-The_extended_phenotype" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_extended_phenotype"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>The extended phenotype</span> </div> </a> <ul id="toc-The_extended_phenotype-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genes_and_phenotypes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Genes_and_phenotypes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Genes and phenotypes</span> </div> </a> <ul id="toc-Genes_and_phenotypes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phenome_and_phenomics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Phenome_and_phenomics"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Phenome and phenomics</span> </div> </a> <ul id="toc-Phenome_and_phenomics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Large-scale_phenotyping_and_genetic_screens" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Large-scale_phenotyping_and_genetic_screens"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Large-scale phenotyping and genetic screens</span> </div> </a> <ul id="toc-Large-scale_phenotyping_and_genetic_screens-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolutionary_origin_of_phenotype" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolutionary_origin_of_phenotype"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Evolutionary origin of phenotype</span> </div> </a> <ul id="toc-Evolutionary_origin_of_phenotype-sublist" class="vector-toc-list"> </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">7</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">8</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">9</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">Phenotype</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 75 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-75" 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">75 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Fenotipe" title="Fenotipe – Afrikaans" lang="af" hreflang="af" data-title="Fenotipe" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%86%D9%85%D8%B7_%D8%B8%D8%A7%D9%87%D8%B1%D9%8A" title="نمط ظاهري – Arabic" lang="ar" hreflang="ar" data-title="نمط ظاهري" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Fenotipu" title="Fenotipu – Asturian" lang="ast" hreflang="ast" data-title="Fenotipu" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Fenotip" title="Fenotip – Azerbaijani" lang="az" hreflang="az" data-title="Fenotip" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/Pi%C3%A1u-hi%C4%81n-h%C3%AAng" title="Piáu-hiān-hêng – Minnan" lang="nan" hreflang="nan" data-title="Piáu-hiān-hêng" 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-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Bulgarian" lang="bg" hreflang="bg" data-title="Фенотип" 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/Fenotip" title="Fenotip – Bosnian" lang="bs" hreflang="bs" data-title="Fenotip" 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/Fenotip" title="Fenotip – Catalan" lang="ca" hreflang="ca" data-title="Fenotip" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Chuvash" lang="cv" hreflang="cv" data-title="Фенотип" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target"><span>Чӑвашла</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Fenotyp" title="Fenotyp – Czech" lang="cs" hreflang="cs" data-title="Fenotyp" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/F%C3%A6notype" title="Fænotype – Danish" lang="da" hreflang="da" data-title="Fænotype" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Ph%C3%A4notyp" title="Phänotyp – German" lang="de" hreflang="de" data-title="Phänotyp" 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/Fenot%C3%BC%C3%BCp" title="Fenotüüp – Estonian" lang="et" hreflang="et" data-title="Fenotüüp" 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%A6%CE%B1%CE%B9%CE%BD%CF%8C%CF%84%CF%85%CF%80%CE%BF%CF%82" title="Φαινότυπος – Greek" lang="el" hreflang="el" data-title="Φαινότυπος" 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/Fenotipo" title="Fenotipo – Spanish" lang="es" hreflang="es" data-title="Fenotipo" 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/Fenotipo" title="Fenotipo – Esperanto" lang="eo" hreflang="eo" data-title="Fenotipo" 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/Fenotipo" title="Fenotipo – Basque" lang="eu" hreflang="eu" data-title="Fenotipo" 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/%D9%81%D9%86%D9%88%D8%AA%DB%8C%D9%BE" title="فنوتیپ – Persian" lang="fa" hreflang="fa" data-title="فنوتیپ" 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/Ph%C3%A9notype" title="Phénotype – French" lang="fr" hreflang="fr" data-title="Phénotype" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Feinit%C3%ADopa" title="Feinitíopa – Irish" lang="ga" hreflang="ga" data-title="Feinitíopa" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Fenotipo" title="Fenotipo – Galician" lang="gl" hreflang="gl" data-title="Fenotipo" 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/%ED%91%9C%ED%98%84%ED%98%95" title="표현형 – Korean" lang="ko" hreflang="ko" data-title="표현형" 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%96%D5%A5%D5%B6%D5%B8%D5%BF%D5%AB%D5%BA" title="Ֆենոտիպ – Armenian" lang="hy" hreflang="hy" data-title="Ֆենոտիպ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Fenotip" title="Fenotip – Croatian" lang="hr" hreflang="hr" data-title="Fenotip" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Fenotipo" title="Fenotipo – Ido" lang="io" hreflang="io" data-title="Fenotipo" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Fenotipe" title="Fenotipe – Indonesian" lang="id" hreflang="id" data-title="Fenotipe" 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/Svipfar" title="Svipfar – Icelandic" lang="is" hreflang="is" data-title="Svipfar" 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/Fenotipo" title="Fenotipo – Italian" lang="it" hreflang="it" data-title="Fenotipo" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he badge-Q70894304 mw-list-item" title=""><a href="https://he.wikipedia.org/wiki/%D7%A4%D7%A0%D7%95%D7%98%D7%99%D7%A4" title="פנוטיפ – Hebrew" lang="he" hreflang="he" data-title="פנוטיפ" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A4%E1%83%94%E1%83%9C%E1%83%9D%E1%83%A2%E1%83%98%E1%83%9E%E1%83%98" title="ფენოტიპი – Georgian" lang="ka" hreflang="ka" data-title="ფენოტიპი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Kazakh" lang="kk" hreflang="kk" data-title="Фенотип" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Fenotip" title="Fenotip – Haitian Creole" lang="ht" hreflang="ht" data-title="Fenotip" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Fenot%C3%AEp" title="Fenotîp – Kurdish" lang="ku" hreflang="ku" data-title="Fenotîp" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Kyrgyz" lang="ky" hreflang="ky" data-title="Фенотип" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Fenotips" title="Fenotips – Latvian" lang="lv" hreflang="lv" data-title="Fenotips" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Phenotyp" title="Phenotyp – Luxembourgish" lang="lb" hreflang="lb" data-title="Phenotyp" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target"><span>Lëtzebuergesch</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Fenot%C3%ADpus" title="Fenotípus – Hungarian" lang="hu" hreflang="hu" data-title="Fenotípus" 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%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Macedonian" lang="mk" hreflang="mk" data-title="Фенотип" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Fenotip" title="Fenotip – Malay" lang="ms" hreflang="ms" data-title="Fenotip" 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%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Mongolian" lang="mn" hreflang="mn" data-title="Фенотип" 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/Fenotype" title="Fenotype – Dutch" lang="nl" hreflang="nl" data-title="Fenotype" 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/%E8%A1%A8%E7%8F%BE%E5%9E%8B" title="表現型 – Japanese" lang="ja" hreflang="ja" data-title="表現型" 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-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Fenotype" title="Fenotype – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Fenotype" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Fenotipe" title="Fenotipe – Occitan" lang="oc" hreflang="oc" data-title="Fenotipe" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Fenotip" title="Fenotip – Uzbek" lang="uz" hreflang="uz" data-title="Fenotip" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%81%DB%90%D9%86%D9%88%D9%BC%D8%A7%DB%8C%D9%BE" title="فېنوټایپ – Pashto" lang="ps" hreflang="ps" data-title="فېنوټایپ" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Ph%C3%A4notyyp" title="Phänotyyp – Low German" lang="nds" hreflang="nds" data-title="Phänotyyp" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target"><span>Plattdüütsch</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Fenotyp" title="Fenotyp – Polish" lang="pl" hreflang="pl" data-title="Fenotyp" 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/Fen%C3%B3tipo" title="Fenótipo – Portuguese" lang="pt" hreflang="pt" data-title="Fenótipo" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-kaa mw-list-item"><a href="https://kaa.wikipedia.org/wiki/Fenotip" title="Fenotip – Kara-Kalpak" lang="kaa" hreflang="kaa" data-title="Fenotip" data-language-autonym="Qaraqalpaqsha" data-language-local-name="Kara-Kalpak" class="interlanguage-link-target"><span>Qaraqalpaqsha</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Fenotip" title="Fenotip – Romanian" lang="ro" hreflang="ro" data-title="Fenotip" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-rue mw-list-item"><a href="https://rue.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Rusyn" lang="rue" hreflang="rue" data-title="Фенотип" data-language-autonym="Русиньскый" data-language-local-name="Rusyn" class="interlanguage-link-target"><span>Русиньскый</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Russian" lang="ru" hreflang="ru" data-title="Фенотип" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Phenoteep" title="Phenoteep – Scots" lang="sco" hreflang="sco" data-title="Phenoteep" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target"><span>Scots</span></a></li><li class="interlanguage-link interwiki-stq mw-list-item"><a href="https://stq.wikipedia.org/wiki/Ph%C3%A4notyp" title="Phänotyp – Saterland Frisian" lang="stq" hreflang="stq" data-title="Phänotyp" data-language-autonym="Seeltersk" data-language-local-name="Saterland Frisian" class="interlanguage-link-target"><span>Seeltersk</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Phenotype" title="Phenotype – Simple English" lang="en-simple" hreflang="en-simple" data-title="Phenotype" 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-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Fenotyp" title="Fenotyp – Slovak" lang="sk" hreflang="sk" data-title="Fenotyp" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Fenotip" title="Fenotip – Slovenian" lang="sl" hreflang="sl" data-title="Fenotip" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%DA%95%D9%88%D9%88%D8%AE%D8%B3%D8%A7%D8%B1%DB%95%D8%A8%D8%A7%D8%A8%DB%95%D8%AA" title="ڕووخسارەبابەت – Central Kurdish" lang="ckb" hreflang="ckb" data-title="ڕووخسارەبابەت" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Serbian" lang="sr" hreflang="sr" data-title="Фенотип" 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/Fenotip" title="Fenotip – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Fenotip" 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/Fenotyyppi" title="Fenotyyppi – Finnish" lang="fi" hreflang="fi" data-title="Fenotyyppi" 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/Fenotyp" title="Fenotyp – Swedish" lang="sv" hreflang="sv" data-title="Fenotyp" 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/Penotipo" title="Penotipo – Tagalog" lang="tl" hreflang="tl" data-title="Penotipo" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%A4%E0%AF%8B%E0%AE%B1%E0%AF%8D%E0%AE%B1%E0%AE%B5%E0%AE%AE%E0%AF%88%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AF%81" title="தோற்றவமைப்பு – Tamil" lang="ta" hreflang="ta" data-title="தோற்றவமைப்பு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-kab mw-list-item"><a href="https://kab.wikipedia.org/wiki/Tumanawt" title="Tumanawt – Kabyle" lang="kab" hreflang="kab" data-title="Tumanawt" data-language-autonym="Taqbaylit" data-language-local-name="Kabyle" class="interlanguage-link-target"><span>Taqbaylit</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A3%E0%B8%B9%E0%B8%9B%E0%B9%81%E0%B8%9A%E0%B8%9A%E0%B8%9B%E0%B8%A3%E0%B8%B2%E0%B8%81%E0%B8%8F" title="รูปแบบปรากฏ – Thai" lang="th" hreflang="th" data-title="รูปแบบปรากฏ" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tg mw-list-item"><a href="https://tg.wikipedia.org/wiki/%D0%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Tajik" lang="tg" hreflang="tg" data-title="Фенотип" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target"><span>Тоҷикӣ</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Fenotip" title="Fenotip – Turkish" lang="tr" hreflang="tr" data-title="Fenotip" 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%A4%D0%B5%D0%BD%D0%BE%D1%82%D0%B8%D0%BF" title="Фенотип – Ukrainian" lang="uk" hreflang="uk" data-title="Фенотип" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B7%D8%B1%D8%B2_%D8%B8%D8%A7%DB%81%D8%B1%DB%8C" title="طرز ظاہری – Urdu" lang="ur" hreflang="ur" data-title="طرز ظاہری" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Ki%E1%BB%83u_h%C3%ACnh" title="Kiểu hình – Vietnamese" lang="vi" hreflang="vi" data-title="Kiểu hình" 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-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E8%A1%A8%E5%9E%8B" title="表型 – 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searchaux" style="display:none">Composite of the organism's observable characteristics or traits</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">For a non-technical introduction to the topic, see <a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction to genetics</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For other uses, see <a href="/wiki/Phenotype_(disambiguation)" class="mw-disambig" title="Phenotype (disambiguation)">Phenotype (disambiguation)</a>.</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Coquina_variation3.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Coquina_variation3.jpg/250px-Coquina_variation3.jpg" decoding="async" width="250" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Coquina_variation3.jpg/375px-Coquina_variation3.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Coquina_variation3.jpg/500px-Coquina_variation3.jpg 2x" data-file-width="3000" data-file-height="3000" /></a><figcaption>The <a href="/wiki/Mollusc_shell" title="Mollusc shell">shells</a> of individuals within the <a href="/wiki/Bivalve" class="mw-redirect" title="Bivalve">bivalve</a> <a href="/wiki/Mollusca" title="Mollusca">mollusk</a> species <i><a href="/wiki/Donax_variabilis" title="Donax variabilis">Donax variabilis</a></i> show diverse <a href="/wiki/Animal_coloration" title="Animal coloration">coloration</a> and <a href="/wiki/Patterns_in_nature" title="Patterns in nature">patterning</a> in their phenotypes.</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Punnett_square_mendel_flowers.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Punnett_square_mendel_flowers.svg/220px-Punnett_square_mendel_flowers.svg.png" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Punnett_square_mendel_flowers.svg/330px-Punnett_square_mendel_flowers.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/17/Punnett_square_mendel_flowers.svg/440px-Punnett_square_mendel_flowers.svg.png 2x" data-file-width="550" data-file-height="550" /></a><figcaption>Here the relation between <a href="/wiki/Genotype" title="Genotype">genotype</a> and phenotype is illustrated, using a <a href="/wiki/Punnett_square" title="Punnett square">Punnett square</a>, for the character of petal color in pea plants. The letters B and b represent <a href="/wiki/Gene" title="Gene">genes</a> for color, and the pictures show the resultant phenotypes. This shows how multiple genotypes (BB and Bb) may yield the same phenotype (purple petals).</figcaption></figure> <p>In <a href="/wiki/Genetics" title="Genetics">genetics</a>, the <b>phenotype</b> (from <a href="/wiki/Ancient_Greek_language" class="mw-redirect" title="Ancient Greek language">Ancient Greek</a> <i> </i><span lang="grc"><a href="https://en.wiktionary.org/wiki/%CF%86%CE%B1%CE%AF%CE%BD%CF%89#Ancient_Greek" class="extiw" title="wikt:φαίνω">φαίνω</a></span><i> (<span title="Ancient Greek transliteration" lang="grc-Latn"><i>phaínō</i></span>)</i> 'to appear, show' and <i> </i><span lang="grc"><a href="https://en.wiktionary.org/wiki/%CF%84%CF%8D%CF%80%CE%BF%CF%82#Ancient_Greek" class="extiw" title="wikt:τύπος">τύπος</a></span><i> (<span title="Ancient Greek transliteration" lang="grc-Latn"><i>túpos</i></span>)</i> 'mark, type') is the set of observable characteristics or <a href="/wiki/Phenotypic_trait" title="Phenotypic trait">traits</a> of an <a href="/wiki/Organism" title="Organism">organism</a>.<sup id="cite_ref-Oxford_Advanced_Learners_Dictionary_at_OxfordLearnersDictionaries.com_1-0" class="reference"><a href="#cite_note-Oxford_Advanced_Learners_Dictionary_at_OxfordLearnersDictionaries.com-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Understanding_Evolution_2-0" class="reference"><a href="#cite_note-Understanding_Evolution-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The term covers the organism's <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a> (physical form and structure), its <a href="/wiki/Developmental_biology" title="Developmental biology">developmental</a> processes, its biochemical and physiological properties, its <a href="/wiki/Behavior" title="Behavior">behavior</a>, and the products of behavior. An organism's phenotype results from two basic factors: the <a href="/wiki/Gene_expression" title="Gene expression">expression</a> of an organism's genetic code (its <a href="/wiki/Genotype" title="Genotype">genotype</a>) and the influence of environmental factors. Both factors may interact, further affecting the phenotype. When two or more clearly different phenotypes exist in the same population of a species, the species is called <a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">polymorphic</a>. A well-documented example of polymorphism is <a href="/wiki/Labrador_Retriever_coat_colour_genetics" title="Labrador Retriever coat colour genetics">Labrador Retriever coloring</a>; while the coat color depends on many genes, it is clearly seen in the environment as yellow, black, and brown. <a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a> in 1978<sup id="cite_ref-r1_3-0" class="reference"><a href="#cite_note-r1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> and then again in his 1982 book <i><a href="/wiki/The_Extended_Phenotype" title="The Extended Phenotype">The Extended Phenotype</a></i> suggested that one can regard <a href="/wiki/Bird_nest" title="Bird nest">bird nests</a> and other built structures such as <a href="/wiki/Caddisfly" title="Caddisfly">caddisfly</a> larva cases and <a href="/wiki/Beaver_dam" title="Beaver dam">beaver dams</a> as "extended phenotypes". </p><p><a href="/wiki/Wilhelm_Johannsen" title="Wilhelm Johannsen">Wilhelm Johannsen</a> proposed the <a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">genotype–phenotype distinction</a> in 1911 to make clear the difference between an organism's <a href="/wiki/Heredity" title="Heredity">hereditary material</a> and what that hereditary material produces.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The distinction resembles that proposed by <a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a> (1834–1914), who distinguished between <a href="/wiki/Germ_plasm" title="Germ plasm">germ plasm</a> (heredity) and <a href="/wiki/Somatic_cell" title="Somatic cell">somatic cells</a> (the body). More recently, in <i><a href="/wiki/The_Selfish_Gene" title="The Selfish Gene">The Selfish Gene</a></i> (1976), Dawkins distinguished these concepts as replicators and vehicles. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Definition">Definition</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=1" title="Edit section: Definition"><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">"Phenome" redirects here. For the speech unit, see <a href="/wiki/Phoneme" title="Phoneme">Phoneme</a>.</div> <p><span class="anchor" id="Definition"></span>Despite its seemingly straightforward definition, the concept of the phenotype has hidden subtleties. It may seem that anything dependent on the <a href="/wiki/Genotype" title="Genotype">genotype</a> is a phenotype, including <a href="/wiki/Molecule" title="Molecule">molecules</a> such as <a href="/wiki/RNA" title="RNA">RNA</a> and <a href="/wiki/Protein" title="Protein">proteins</a>. Most molecules and structures coded by the genetic material are not visible in the appearance of an organism, yet they are observable (for example by <a href="/wiki/Western_blot" title="Western blot">Western blotting</a>) and are thus part of the phenotype; human <a href="/wiki/Human_blood_group_systems" title="Human blood group systems">blood groups</a> are an example. It may seem that this goes beyond the original intentions of the concept with its focus on the (living) organism in itself. Either way, the term phenotype includes inherent traits or characteristics that are observable or traits that can be made visible by some technical procedure.<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. (May 2024)">citation needed</span></a></i>]</sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:ABO_Blood_Group_Phenotypes.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/ABO_Blood_Group_Phenotypes.jpg/220px-ABO_Blood_Group_Phenotypes.jpg" decoding="async" width="220" height="272" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/ABO_Blood_Group_Phenotypes.jpg/330px-ABO_Blood_Group_Phenotypes.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/ABO_Blood_Group_Phenotypes.jpg/440px-ABO_Blood_Group_Phenotypes.jpg 2x" data-file-width="1393" data-file-height="1722" /></a><figcaption>ABO blood groups determined through a Punnett square and displaying phenotypes and genotypes</figcaption></figure> <p>The term "phenotype" has sometimes been incorrectly used as a shorthand for the phenotypic difference between a mutant and its <a href="/wiki/Wild_type" title="Wild type">wild type</a>, which would lead to the false statement that a "mutation has no phenotype".<sup id="cite_ref-pmid12884976_6-0" class="reference"><a href="#cite_note-pmid12884976-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Behaviors and their consequences are also phenotypes, since behaviors are observable characteristics. <b>Behavioral phenotypes</b> include cognitive, personality, and behavioral patterns. Some behavioral phenotypes may characterize psychiatric disorders<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> or syndromes.<sup id="cite_ref-O'Brien&Yule19952_8-0" class="reference"><a href="#cite_note-O'Brien&Yule19952-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-O'Brien20022_9-0" class="reference"><a href="#cite_note-O'Brien20022-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>A <b>phenome</b> is the set of all traits expressed by a <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>, <a href="/wiki/Biological_tissue" class="mw-redirect" title="Biological tissue">tissue</a>, <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organ</a>, <a href="/wiki/Organism" title="Organism">organism</a>, or <a href="/wiki/Species" title="Species">species</a>. The term was first used by Davis in 1949, "We here propose the name <i>phenome</i> for the sum total of extragenic, non-autoreproductive portions of the cell, whether cytoplasmic or nuclear. The phenome would be the material basis of the phenotype, just as the genome is the material basis of the <a href="/wiki/Genotype" title="Genotype">genotype</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> </p><p>Although phenome has been in use for many years, the distinction between the use of phenome and phenotype is problematic. A proposed definition for both terms as the "physical totality of all traits of an organism or of one of its subsystems" was put forth by Mahner and Kary in 1997, who argue that although scientists tend to intuitively use these and related terms in a manner that does not impede research, the terms are not well defined and usage of the terms is not consistent.<sup id="cite_ref-JTB1_11-0" class="reference"><a href="#cite_note-JTB1-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Some usages of the term suggest that the phenome of a given organism is best understood as a kind of matrix of data representing physical manifestation of phenotype. For example, discussions led by A. Varki among those who had used the term up to 2003 suggested the following definition: "The body of information describing an organism's phenotypes, under the influences of genetic and environmental factors".<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> Another team of researchers characterize "the human phenome [as] a multidimensional search space with several neurobiological levels, spanning the proteome, cellular systems (e.g., signaling pathways), neural systems and cognitive and behavioural phenotypes."<sup id="cite_ref-Neuroscience1_13-0" class="reference"><a href="#cite_note-Neuroscience1-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Plant biologists have started to explore the phenome in the study of plant physiology.<sup id="cite_ref-tplants_14-0" class="reference"><a href="#cite_note-tplants-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>In 2009, a research team demonstrated the feasibility of identifying genotype–phenotype associations using <a href="/wiki/Electronic_health_records" class="mw-redirect" title="Electronic health records">electronic health records</a> (EHRs) linked to DNA <a href="/wiki/Biobanks" class="mw-redirect" title="Biobanks">biobanks</a>. They called this method <a href="/wiki/Phenome-wide_association_study" title="Phenome-wide association study">phenome-wide association study</a> (PheWAS).<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Pan_Y_G_E.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/Pan_Y_G_E.jpg/384px-Pan_Y_G_E.jpg" decoding="async" width="384" height="221" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/Pan_Y_G_E.jpg/576px-Pan_Y_G_E.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/13/Pan_Y_G_E.jpg/768px-Pan_Y_G_E.jpg 2x" data-file-width="1956" data-file-height="1125" /></a><figcaption>Exploring relationships among phenotype, genotype and environment at different levels<sup id="cite_ref-:0_16-0" class="reference"><a href="#cite_note-:0-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Inspired by the evolution from genotype to genome to <a href="/wiki/Pan-genome" title="Pan-genome">pan-genome</a>, a concept of exploring the relationship ultimately among pan-phenome, <a href="/wiki/Pan-genome" title="Pan-genome">pan-genome</a>, and pan-<a href="/wiki/Envirome" title="Envirome">envirome</a> was proposed in 2023.<sup id="cite_ref-:0_16-1" class="reference"><a href="#cite_note-:0-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Biston.betularia.7200.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/220px-Biston.betularia.7200.jpg" decoding="async" width="220" height="131" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/330px-Biston.betularia.7200.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Biston.betularia.7200.jpg/440px-Biston.betularia.7200.jpg 2x" data-file-width="2798" data-file-height="1662" /></a><figcaption><i><a href="/wiki/Biston_betularia" class="mw-redirect" title="Biston betularia">Biston betularia</a></i> morpha <i>typica</i>, the standard light-colored peppered moth</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Biston.betularia.f.carbonaria.7209.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/220px-Biston.betularia.f.carbonaria.7209.jpg" decoding="async" width="220" height="107" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/330px-Biston.betularia.f.carbonaria.7209.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Biston.betularia.f.carbonaria.7209.jpg/440px-Biston.betularia.f.carbonaria.7209.jpg 2x" data-file-width="2816" data-file-height="1366" /></a><figcaption><i>B.betularia</i> morpha <i>carbonaria</i>, the melanic form, illustrating discontinuous variation</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Phenotypic_variation">Phenotypic variation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=2" title="Edit section: Phenotypic variation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Ecophenotypic_variation" title="Ecophenotypic variation">Ecophenotypic variation</a></div> <p>Phenotypic variation (due to underlying heritable <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic variation</a>) is a fundamental prerequisite for <a href="/wiki/Evolution" title="Evolution">evolution</a> by <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>. It is the living organism as a whole that contributes (or not) to the next generation, so natural selection affects the genetic structure of a population indirectly via the contribution of phenotypes. Without phenotypic variation, there would be no evolution by natural selection.<sup id="cite_ref-Lewontin70_17-0" class="reference"><a href="#cite_note-Lewontin70-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>The interaction between genotype and phenotype has often been conceptualized by the following relationship: </p> <dl><dd>genotype (G) + environment (E) → phenotype (P)</dd></dl> <p>A more nuanced version of the relationship is: </p> <dl><dd>genotype (G) + environment (E) + genotype & environment interactions (GE) → phenotype (P)</dd></dl> <p>Genotypes often have much flexibility in the modification and expression of phenotypes; in many organisms these phenotypes are very different under varying environmental conditions. The plant <i><a href="/wiki/Hieracium_umbellatum" title="Hieracium umbellatum">Hieracium umbellatum</a></i> is found growing in two different <a href="/wiki/Habitat" title="Habitat">habitats</a> in Sweden. One habitat is rocky, sea-side <a href="/wiki/Cliff" title="Cliff">cliffs</a>, where the plants are bushy with broad leaves and expanded <a href="/wiki/Inflorescence" title="Inflorescence">inflorescences</a>; the other is among <a href="/wiki/Dune" title="Dune">sand dunes</a> where the plants grow prostrate with narrow leaves and compact inflorescences. These habitats alternate along the coast of Sweden and the habitat that the seeds of <i>Hieracium umbellatum</i> land in, determine the phenotype that grows.<sup id="cite_ref-Botanyonline_18-0" class="reference"><a href="#cite_note-Botanyonline-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>An example of random variation in <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> flies is the number of <a href="/wiki/Ommatidia" class="mw-redirect" title="Ommatidia">ommatidia</a>, which may vary (randomly) between left and right eyes in a single individual as much as they do between different genotypes overall, or between <a href="/wiki/Cloning" title="Cloning">clones</a> raised in different environments.<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. (March 2018)">citation needed</span></a></i>]</sup> </p><p>The concept of phenotype can be extended to variations below the level of the <a href="/wiki/Gene" title="Gene">gene</a> that affect an organism's fitness. For example, <a href="/wiki/Silent_mutations" class="mw-redirect" title="Silent mutations">silent mutations</a> that do not change the corresponding amino acid sequence of a gene may change the frequency of <a href="/wiki/Guanine" title="Guanine">guanine</a>-<a href="/wiki/Cytosine" title="Cytosine">cytosine</a> base pairs (<a href="/wiki/GC_content" class="mw-redirect" title="GC content">GC content</a>). These base pairs have a higher thermal stability (<i>melting point</i>) than <a href="/wiki/Adenine" title="Adenine">adenine</a>-<a href="/wiki/Thymine" title="Thymine">thymine</a>, a property that might convey, among organisms living in high-temperature environments, a selective advantage on variants enriched in GC content.<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. (March 2018)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="The_extended_phenotype">The extended phenotype</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=3" title="Edit section: The extended phenotype"><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/The_Extended_Phenotype" title="The Extended Phenotype">The Extended Phenotype</a></div> <p><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a> described a phenotype that included all effects that a gene has on its surroundings, including other organisms, as an extended phenotype, arguing that "An animal's behavior tends to maximize the survival of the genes 'for' that behavior, whether or not those genes happen to be in the body of the particular animal performing it."<sup id="cite_ref-r1_3-1" class="reference"><a href="#cite_note-r1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> For instance, an organism such as a <a href="/wiki/Beaver" title="Beaver">beaver</a> modifies its environment by building a <a href="/wiki/Beaver_dam" title="Beaver dam">beaver dam</a>; this can be considered an <a href="/wiki/Gene_expression" title="Gene expression">expression of its genes</a>, just as its <a href="/wiki/Incisor" title="Incisor">incisor</a> teeth are—which it uses to modify its environment. Similarly, when a bird feeds a <a href="/wiki/Brood_parasite" class="mw-redirect" title="Brood parasite">brood parasite</a> such as a <a href="/wiki/Cuckoo" title="Cuckoo">cuckoo</a>, it is unwittingly extending its phenotype; and when genes in an <a href="/wiki/Orchid" title="Orchid">orchid</a> affect <a href="/wiki/Orchid_bee" class="mw-redirect" title="Orchid bee">orchid bee</a> behavior to increase pollination, or when genes in a <a href="/wiki/Peacock" class="mw-redirect" title="Peacock">peacock</a> affect the copulatory decisions of peahens, again, the phenotype is being extended. Genes are, in Dawkins's view, selected by their phenotypic effects.<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><p>Other biologists broadly agree that the extended phenotype concept is relevant, but consider that its role is largely explanatory, rather than assisting in the design of experimental tests.<sup id="cite_ref-Hunter2009_20-0" class="reference"><a href="#cite_note-Hunter2009-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genes_and_phenotypes">Genes and phenotypes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=4" title="Edit section: Genes and phenotypes"><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:Genotype_to_phenotype.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Genotype_to_phenotype.svg/220px-Genotype_to_phenotype.svg.png" decoding="async" width="220" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Genotype_to_phenotype.svg/330px-Genotype_to_phenotype.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/65/Genotype_to_phenotype.svg/440px-Genotype_to_phenotype.svg.png 2x" data-file-width="395" data-file-height="244" /></a><figcaption>An organism's phenotype is determined by the sum of its genetic material along with the influence of its environment. This is mediated by a range of biological mechanisms: either the direct activities of gene products or their downstream effects.<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></figcaption></figure> <p>Phenotypes are determined by an interaction of genes and the environment, but the mechanism for each gene and phenotype is different. For instance, an <a href="/wiki/Albinism" title="Albinism">albino</a> phenotype may be caused by a mutation in the gene encoding <a href="/wiki/Tyrosinase" title="Tyrosinase">tyrosinase</a> which is a key enzyme in <a href="/wiki/Melanin" title="Melanin">melanin</a> formation. However, exposure to <a href="/wiki/Ultraviolet" title="Ultraviolet">UV radiation</a> can increase melanin production, hence the environment plays a role in this phenotype as well. For most complex phenotypes the precise genetic mechanism remains unknown. For instance, it is largely unclear how genes determine the shape of bones or the human ear.<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. (March 2023)">citation needed</span></a></i>]</sup> </p><p>Gene expression plays a crucial role in determining the phenotypes of organisms. The level of gene expression can affect the phenotype of an organism. For example, if a gene that codes for a particular <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> is expressed at high levels, the organism may produce more of that enzyme and exhibit a particular trait as a result. On the other hand, if the gene is expressed at low levels, the organism may produce less of the enzyme and exhibit a different trait.<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>Gene expression is regulated at various levels and thus each level can affect certain phenotypes, including <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcriptional</a> and post-transcriptional regulation.<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. (May 2024)">citation needed</span></a></i>]</sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Tortie-flame.jpg" class="mw-file-description"><img alt="tortoiseshell cat" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Tortie-flame.jpg/220px-Tortie-flame.jpg" decoding="async" width="220" height="187" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Tortie-flame.jpg/330px-Tortie-flame.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/3/32/Tortie-flame.jpg 2x" data-file-width="383" data-file-height="326" /></a><figcaption>The patchy colors of a tortoiseshell cat are the result of different levels of expression of pigmentation genes in different areas of the skin.</figcaption></figure> <p>Changes in the levels of gene expression can be influenced by a variety of factors, such as environmental conditions, genetic variations, and <a href="/wiki/Epigenetics" title="Epigenetics">epigenetic</a> modifications. These modifications can be influenced by environmental factors such as diet, stress, and exposure to toxins, and can have a significant impact on an individual's phenotype. Some phenotypes may be the result of changes in gene expression due to these factors, rather than changes in genotype. An experiment involving <a href="/wiki/Machine_learning" title="Machine learning">machine learning</a> methods utilizing gene expressions measured from RNA sequencing found that they can contain enough signal to separate individuals in the context of phenotype prediction.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Phenome_and_phenomics">Phenome and phenomics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=5" title="Edit section: Phenome and phenomics"><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">Not to be confused with <a href="/wiki/Phoneme" title="Phoneme">Phoneme</a> or <a href="/wiki/Phonology" title="Phonology">Phonology</a>.</div> <p>Although a phenotype is the ensemble of observable characteristics displayed by an organism, the word <i><a href="/wiki/Phenome" class="mw-redirect" title="Phenome">phenome</a></i> is sometimes used to refer to a collection of traits, while the simultaneous study of such a collection is referred to as <i><a href="/wiki/Phenomics" title="Phenomics">phenomics</a></i>.<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><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> Phenomics is an important field of study because it can be used to figure out which genomic variants affect phenotypes which then can be used to explain things like health, disease, and evolutionary fitness.<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> Phenomics forms a large part of the <a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a>.<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>Phenomics has applications in agriculture. For instance, genomic variations such as drought and heat resistance can be identified through phenomics to create more durable GMOs.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-tplants_14-1" class="reference"><a href="#cite_note-tplants-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>Phenomics may be a stepping stone towards <a href="/wiki/Personalized_medicine" title="Personalized medicine">personalized medicine</a>, particularly <a href="/wiki/Drug_therapy" class="mw-redirect" title="Drug therapy">drug therapy</a>.<sup id="cite_ref-monte_29-0" class="reference"><a href="#cite_note-monte-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> Once the phenomic database has acquired enough data, a person's phenomic information can be used to select specific drugs tailored to the individual.<sup id="cite_ref-monte_29-1" class="reference"><a href="#cite_note-monte-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Large-scale_phenotyping_and_genetic_screens">Large-scale phenotyping and genetic screens</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=6" title="Edit section: Large-scale phenotyping and genetic screens"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Genetic_screen" title="Genetic screen">Genetic screen</a> and <a href="/wiki/Essential_gene" title="Essential gene">Essential gene</a></div> <p>Large-scale genetic screens can identify the genes or <a href="/wiki/Mutation" title="Mutation">mutations</a> that affect the phenotype of an organism. Analyzing the phenotypes of mutant genes can also aid in determining gene function.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Most genetic screens have used microorganisms, in which genes can be easily deleted. For instance, nearly all genes have been deleted in <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a></i><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> and many other <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, but also in several eukaryotic model organisms such as <a href="/wiki/Baker%27s_yeast" title="Baker's yeast">baker's yeast</a><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Schizosaccharomyces_pombe" title="Schizosaccharomyces pombe">fission yeast</a>.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Among other discoveries, such studies have revealed lists of essential genes . </p><p>More recently, large-scale <a href="/wiki/Phenotypic_screening" title="Phenotypic screening">phenotypic screens</a> have also been used in animals, e.g. to study lesser understood phenotypes such as <a href="/wiki/Behavioural_genetics" title="Behavioural genetics">behavior</a>. In one screen, the role of mutations in mice were studied in areas such as learning and <a href="/wiki/Memory" title="Memory">memory</a>, <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">circadian rhythmicity</a>, vision, responses to stress and response to <a href="/wiki/Stimulant" title="Stimulant">psychostimulants</a>. </p> <table class="wikitable"> <caption><a href="//doi.org/10.1016/j.tins.2006.02.006" class="extiw" title="doi:10.1016/j.tins.2006.02.006">Large-scale mutagenesis and phenotypic screens for the nervous system and behavior in mice</a> </caption> <tbody><tr> <th>Phenotypic Domain </th> <th>Assay </th> <th>Notes </th> <th>Software Package </th></tr> <tr> <td>Circadian Rhythm </td> <td>Wheel running behavior </td> <td> </td> <td>ClockLab </td></tr> <tr> <td>Learning and Memory </td> <td><a href="/wiki/Fear_conditioning" title="Fear conditioning">Fear conditioning</a> </td> <td>Video-image-based scoring of freezing </td> <td>FreezeFrame </td></tr> <tr> <td>Preliminary Assessment </td> <td><a href="/wiki/Open_field_(animal_test)" title="Open field (animal test)">Open field activity</a> and <a href="/wiki/Elevated_plus_maze" title="Elevated plus maze">elevated plus maze</a> </td> <td>Video-image-based scoring of exploration </td> <td>LimeLight </td></tr> <tr> <td>Psychostimulant response </td> <td>Hyperlocomotion behavior </td> <td>Video-image-based tracking of locomotion </td> <td>BigBrother </td></tr> <tr> <td>Vision </td> <td><a href="/wiki/Electroretinography" title="Electroretinography">Electroretinogram</a> and <a href="/wiki/Fundus_photography" title="Fundus photography">Fundus photography</a> </td> <td> </td> <td>L. Pinto and colleagues </td></tr></tbody></table> <p>This experiment involved the progeny of mice treated with <a href="/wiki/ENU" title="ENU">ENU</a>, or N-ethyl-N-nitrosourea, which is a potent mutagen that causes <a href="/wiki/Point_mutation" title="Point mutation">point mutations</a>. The mice were phenotypically screened for alterations in the different behavioral domains in order to find the number of putative mutants (see table for details). Putative mutants are then tested for heritability in order to help determine the inheritance pattern as well as map out the mutations. Once they have been mapped out, cloned, and identified, it can be determined whether a mutation represents a new gene or not. </p> <table class="wikitable"> <tbody><tr> <th>Phenotypic domain </th> <th>ENU Progeny screened </th> <th>Putative mutants </th> <th>Putative mutant lines with progeny </th> <th>Confirmed mutants </th></tr> <tr> <td>General assessment </td> <td>29860 </td> <td>80 </td> <td>38 </td> <td>14 </td></tr> <tr> <td>Learning and memory </td> <td>23123 </td> <td>165 </td> <td>106 </td> <td>19 </td></tr> <tr> <td>Psychostimulant response </td> <td>20997 </td> <td>168 </td> <td>86 </td> <td>9 </td></tr> <tr> <td><a href="/wiki/Neuroendocrinology" title="Neuroendocrinology">Neuroendocrine</a> response to stress </td> <td>13118 </td> <td>126 </td> <td>54 </td> <td>2 </td></tr> <tr> <td>Vision </td> <td>15582 </td> <td>108 </td> <td>60 </td> <td>6 </td></tr></tbody></table> <p>These experiments showed that mutations in the <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a> gene affected vision and can even cause <a href="/w/index.php?title=Retinal_degeneration_(rhodopsin_mutation)&action=edit&redlink=1" class="new" title="Retinal degeneration (rhodopsin mutation) (page does not exist)">retinal degeneration</a> in mice.<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> The same <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> change causes <a href="/wiki/Blindness" class="mw-redirect" title="Blindness">human familial blindness</a>, showing how phenotyping in animals can inform medical diagnostics and possibly therapy. </p> <div class="mw-heading mw-heading2"><h2 id="Evolutionary_origin_of_phenotype">Evolutionary origin of phenotype</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=7" title="Edit section: Evolutionary origin of phenotype"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/RNA_world" title="RNA world">RNA world</a> is the hypothesized pre-cellular stage in the evolutionary history of life on earth, in which self-replicating RNA molecules proliferated prior to the evolution of <a href="/wiki/DNA" title="DNA">DNA</a> and proteins.<sup id="cite_ref-Michod1983_35-0" class="reference"><a href="#cite_note-Michod1983-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> The folded <a href="/wiki/Three-dimensional" class="mw-redirect" title="Three-dimensional">three-dimensional</a> physical structure of the first RNA molecule that possessed ribozyme activity promoting replication while avoiding destruction would have been the first phenotype, and the <a href="/wiki/Nucleic_acid_sequence" title="Nucleic acid sequence">nucleotide sequence</a> of the first self-replicating RNA molecule would have been the original genotype.<sup id="cite_ref-Michod1983_35-1" class="reference"><a href="#cite_note-Michod1983-35"><span class="cite-bracket">[</span>35<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=Phenotype&action=edit&section=8" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></li> <li><a href="/wiki/Ecotype" title="Ecotype">Ecotype</a></li> <li><a href="/wiki/Endophenotype" title="Endophenotype">Endophenotype</a></li> <li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Molecular_phenotyping" title="Molecular phenotyping">Molecular phenotyping</a></li> <li><a href="/wiki/Phenomics" title="Phenomics">Phenomics</a></li> <li><a href="/wiki/Phenotypic_trait" title="Phenotypic trait">Phenotypic trait</a></li> <li><a href="/wiki/Physiome" title="Physiome">Physiome</a></li> <li><a href="/wiki/Physiomics" title="Physiomics">Physiomics</a></li> <li><a href="/wiki/Race_and_genetics" title="Race and genetics">Race and genetics</a></li> <li><a href="/wiki/Systems_biology" title="Systems biology">Systems biology</a></li> <li><a href="/wiki/List_of_omics_topics_in_biology" title="List of omics topics in biology">List of omics topics in biology</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=9" 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-Oxford_Advanced_Learners_Dictionary_at_OxfordLearnersDictionaries.com-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Oxford_Advanced_Learners_Dictionary_at_OxfordLearnersDictionaries.com_1-0">^</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 a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.oxfordlearnersdictionaries.com/definition/english/phenotype?q=phenotype">"Phenotype adjective – Definition, pictures, pronunciation and usage notes"</a>. <i>Oxford Advanced Learner's Dictionary at OxfordLearnersDictionaries.com</i><span class="reference-accessdate">. Retrieved <span class="nowrap">2020-04-29</span></span>. <q>the set of observable characteristics of an individual resulting from the interaction of its genotype with the environment.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Oxford+Advanced+Learner%27s+Dictionary+at+OxfordLearnersDictionaries.com&rft.atitle=Phenotype+adjective+%E2%80%93+Definition%2C+pictures%2C+pronunciation+and+usage+notes&rft_id=https%3A%2F%2Fwww.oxfordlearnersdictionaries.com%2Fdefinition%2Fenglish%2Fphenotype%3Fq%3Dphenotype&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhenotype" class="Z3988"></span></span> </li> <li id="cite_note-Understanding_Evolution-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-Understanding_Evolution_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://evolution.berkeley.edu/evolibrary/article/genovspheno_01">"Genotype versus phenotype"</a>. <i>Understanding Evolution</i><span class="reference-accessdate">. Retrieved <span class="nowrap">2020-04-29</span></span>. <q>An organism's genotype is the set of genes that it carries. 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class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3761413">3761413</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16519954">16519954</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Trends+in+Neurosciences&rft.atitle=Large-scale+mutagenesis+and+phenotypic+screens+for+the+nervous+system+and+behavior+in+mice&rft.volume=29&rft.issue=4&rft.pages=233-240&rft.date=2006-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3761413%23id-name%3DPMC&rft_id=info%3Apmid%2F16519954&rft_id=info%3Adoi%2F10.1016%2Fj.tins.2006.02.006&rft.aulast=Vitaterna&rft.aufirst=MH&rft.au=Pinto%2C+LH&rft.au=Takahashi%2C+JS&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3761413&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhenotype" class="Z3988"></span></span> </li> <li id="cite_note-Michod1983-35"><span class="mw-cite-backlink">^ <a href="#cite_ref-Michod1983_35-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Michod1983_35-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichod1983" class="citation journal cs1">Michod RE (February 1983). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Ficb%2F23.1.5">"Population biology of the first replicators: on the origin of the genotype, phenotype and organism"</a>. <i>American Zoologist</i>. <b>23</b> (1): 5–14. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Ficb%2F23.1.5">10.1093/icb/23.1.5</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Zoologist&rft.atitle=Population+biology+of+the+first+replicators%3A+on+the+origin+of+the+genotype%2C+phenotype+and+organism.&rft.volume=23&rft.issue=1&rft.pages=5-14&rft.date=1983-02&rft_id=info%3Adoi%2F10.1093%2Ficb%2F23.1.5&rft.aulast=Michod&rft.aufirst=RE&rft_id=https%3A%2F%2Fdoi.org%2F10.1093%252Ficb%252F23.1.5&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhenotype" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Phenotype&action=edit&section=10" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Phenotypes" class="extiw" title="commons:Category:Phenotypes">Phenotypes</a></span>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/40px-Wiktionary-logo-en-v2.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/60px-Wiktionary-logo-en-v2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></span></span></div> <div class="side-box-text plainlist">Look up <i><b><a href="https://en.wiktionary.org/wiki/phenotype" class="extiw" title="wiktionary:phenotype"> phenotype</a></b></i> in Wiktionary, the free dictionary.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://www.jax.org/phenome">Mouse Phenome Database</a></li> <li><a rel="nofollow" class="external text" href="http://www.human-phenotype-ontology.org">Human Phenotype Ontology</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070426154232/http://www.europhenome.org/">Europhenome: Access to raw and annotated mouse phenotype data</a></li> <li><a rel="nofollow" class="external text" href="http://embryo.asu.edu/handle/10776/4206/">"Wilhelm Johannsen's Genotype-Phenotype Distinction" by E. Peirson at the Embryo Project Encyclopedia</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20060909075405/http://phenome.jax.org/pub-cgi/phenome/mpdcgi?rtn=docs%2Fhome">Mouse Phenome Project</a> at the <a href="/wiki/Jackson_Laboratory" title="Jackson Laboratory">Jackson Laboratory</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul 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.navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="The_development_of_phenotype" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a class="mw-selflink selflink">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Systems</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a href="/wiki/Morphogen" title="Morphogen">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><i><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a>: National <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q104053#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4248244-6">Germany</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh96012165">United States</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="fenotyp"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph301381&CON_LNG=ENG">Czech Republic</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007546837305171">Israel</a></span></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; 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