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H. Waddington</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 13 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-13" 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">13 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%83%D9%88%D9%86%D8%B1%D8%A7%D8%AF_%D9%87%D8%A7%D9%84_%D9%88%D8%A7%D8%AF%D9%8A%D9%86%D8%AC%D8%AA%D9%88%D9%86" title="كونراد هال وادينجتون – Arabic" lang="ar" hreflang="ar" data-title="كونراد هال وادينجتون" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%DA%A9%D9%86%D8%B1%D8%A7%D8%AF_%D9%87%D9%84_%D9%88%D8%AF%DB%8C%D9%86%D9%82%D8%AA%D9%88%D9%86" title="کنراد هل ودینقتون – South Azerbaijani" lang="azb" hreflang="azb" data-title="کنراد هل ودینقتون" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target"><span>تۆرکجه</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Conrad_Hal_Waddington" title="Conrad Hal Waddington – German" lang="de" hreflang="de" data-title="Conrad Hal Waddington" 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/Conrad_Hal_Waddington" title="Conrad Hal Waddington – Estonian" lang="et" hreflang="et" data-title="Conrad Hal Waddington" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Conrad_Hal_Waddington" title="Conrad Hal Waddington – Spanish" lang="es" hreflang="es" data-title="Conrad Hal Waddington" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%A9%D9%86%D8%B1%D8%A7%D8%AF_%D9%87%D9%84_%D9%88%D8%AF%DB%8C%D9%86%DA%AF%D8%AA%D9%88%D9%86" title="کنراد هل ودینگتون – Persian" lang="fa" hreflang="fa" data-title="کنراد هل ودینگتون" 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/Conrad_Hal_Waddington" title="Conrad Hal Waddington – French" lang="fr" hreflang="fr" data-title="Conrad Hal Waddington" 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/Conrad_Hal_Waddington" title="Conrad Hal Waddington – Irish" lang="ga" hreflang="ga" data-title="Conrad Hal Waddington" 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/C._H._Waddington" title="C. H. Waddington – Galician" lang="gl" hreflang="gl" data-title="C. H. Waddington" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Conrad_Hal_Waddington" title="Conrad Hal Waddington – Italian" lang="it" hreflang="it" data-title="Conrad Hal Waddington" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A3%D0%BE%D0%B4%D0%B4%D0%B8%D0%BD%D0%B3%D1%82%D0%BE%D0%BD,_%D0%9A%D0%BE%D0%BD%D1%80%D0%B0%D0%B4_%D0%A5%D1%8D%D0%BB" title="Уоддингтон, Конрад Хэл – Russian" lang="ru" hreflang="ru" data-title="Уоддингтон, Конрад Хэл" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9A%D0%BE%D0%BD%D1%80%D0%B0%D0%B4_%D0%92%D0%BE%D0%B4%D0%B4%D1%96%D0%BD%D0%B3%D1%82%D0%BE%D0%BD" title="Конрад Воддінгтон – Ukrainian" lang="uk" hreflang="uk" data-title="Конрад Воддінгтон" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%BA%B7%E6%8B%89%E5%BE%B7%C2%B7%E5%93%88%E5%B0%94%C2%B7%E6%B2%83%E4%B8%81%E9%A1%BF" title="康拉德·哈尔·沃丁顿 – Chinese" lang="zh" hreflang="zh" data-title="康拉德·哈尔·沃丁顿" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q721128#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit 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class="mbox-text-span">This article includes a list of <a href="/wiki/Wikipedia:Citing_sources#General_references" title="Wikipedia:Citing sources">general references</a>, but <b>it lacks sufficient corresponding <a href="/wiki/Wikipedia:Citing_sources#Inline_citations" title="Wikipedia:Citing sources">inline citations</a></b>.<span class="hide-when-compact"> Please help to <a href="/wiki/Wikipedia:WikiProject_Reliability" title="Wikipedia:WikiProject Reliability">improve</a> this article by <a href="/wiki/Wikipedia:When_to_cite" title="Wikipedia:When to cite">introducing</a> more precise citations.</span> <span class="date-container"><i>(<span class="date">December 2022</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> </div> </div><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox biography vcard"><tbody><tr><th colspan="2" class="infobox-above" style="font-size:125%;"><div class="fn">Conrad Hal Waddington</div><div class="honorific-suffix" style="font-size: 77%; font-weight: normal;"><span class="nobold noexcerpt nowraplinks" style="font-size:;"><span style="font-size: 100%;"><a href="/wiki/Commander_of_the_Order_of_the_British_Empire" class="mw-redirect" title="Commander of the Order of the British Empire">CBE</a></span> <span style="font-size: 100%;"><a href="/wiki/Fellow_of_the_Royal_Society" title="Fellow of the Royal Society">FRS</a></span> <span style="font-size: 100%;"><a href="/wiki/Fellow_of_the_Royal_Society_of_Edinburgh" class="mw-redirect" title="Fellow of the Royal Society of Edinburgh">FRSE</a></span></span></div></th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Conrad_Hal_Waddington.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/9/9d/Conrad_Hal_Waddington.jpg/220px-Conrad_Hal_Waddington.jpg" decoding="async" width="220" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/9/9d/Conrad_Hal_Waddington.jpg 1.5x" data-file-width="292" data-file-height="332" /></a></span><div class="infobox-caption">Conrad Hal Waddington in 1934</div></td></tr><tr><th scope="row" class="infobox-label">Born</th><td class="infobox-data">8 November 1905<br /><div style="display:inline" class="birthplace"><a href="/wiki/Evesham" title="Evesham">Evesham</a>, <a href="/wiki/Worcestershire" title="Worcestershire">Worcestershire</a>, England</div></td></tr><tr><th scope="row" class="infobox-label">Died</th><td class="infobox-data">26 September 1975<span style="display:none"> (<span class="dtend dday deathdate">1975-09-27</span>)</span> (aged 69)<br /><div style="display:inline" class="deathplace"><a href="/wiki/Edinburgh" title="Edinburgh">Edinburgh</a>, Scotland</div></td></tr><tr><th scope="row" class="infobox-label">Alma mater</th><td class="infobox-data"><a href="/wiki/University_of_Cambridge" title="University of Cambridge">University of Cambridge</a></td></tr><tr><th scope="row" class="infobox-label">Known for</th><td class="infobox-data"><a href="/wiki/Epigenetic_landscape" class="mw-redirect" title="Epigenetic landscape">Epigenetic landscape</a>, <a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">canalisation</a>, <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetic assimilation</a>, <a href="/wiki/Creode" title="Creode">creode</a></td></tr><tr><th scope="row" class="infobox-label">Spouse</th><td class="infobox-data"><a href="/wiki/Justin_Blanco_White" title="Justin Blanco White">Justin Blanco White</a></td></tr><tr><th scope="row" class="infobox-label">Children</th><td class="infobox-data">3, including <a href="/wiki/Caroline_Humphrey" title="Caroline Humphrey">Caroline Humphrey</a> and <a href="/wiki/Dusa_McDuff" title="Dusa McDuff">Dusa McDuff</a></td></tr><tr><th scope="row" class="infobox-label">Awards</th><td class="infobox-data"><a href="/wiki/Mendel_Medal_(genetics)" class="mw-redirect" title="Mendel Medal (genetics)">Mendel Medal</a> <span style="font-size:85%;">(1960)</span></td></tr><tr><td colspan="2" class="infobox-full-data"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><b>Scientific career</b></td></tr><tr><th scope="row" class="infobox-label">Fields</th><td class="infobox-data category"><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a>, <a href="/wiki/Genetics" title="Genetics">genetics</a>, <a href="/wiki/Paleontology" title="Paleontology">paleontology</a></td></tr><tr><th scope="row" class="infobox-label">Institutions</th><td class="infobox-data"><a href="/wiki/University_of_Edinburgh" title="University of Edinburgh">University of Edinburgh</a><br /><a href="/wiki/University_of_Cambridge" title="University of Cambridge">University of Cambridge</a>, <a href="/wiki/Christ%27s_College,_Cambridge" title="Christ's College, Cambridge">Christ's College</a><br /><a href="/wiki/Wesleyan_University" title="Wesleyan University">Wesleyan University</a><br /><a href="/wiki/Centre_for_Human_Ecology" title="Centre for Human Ecology">Centre for Human Ecology</a></td></tr><tr><th scope="row" class="infobox-label">Doctoral students</th><td class="infobox-data"><a href="/wiki/Robert_Edwards_(physiologist)" title="Robert Edwards (physiologist)">Robert Edwards</a></td></tr><tr style="display:none"><td colspan="2"> </td></tr></tbody></table> <p><b>Conrad Hal Waddington</b> <span class="nobold noexcerpt nowraplinks" style="font-size:;"><span style="font-size: 85%;"><a href="/wiki/Commander_of_the_Order_of_the_British_Empire" class="mw-redirect" title="Commander of the Order of the British Empire">CBE</a></span> <span style="font-size: 85%;"><a href="/wiki/Fellow_of_the_Royal_Society" title="Fellow of the Royal Society">FRS</a></span> <span style="font-size: 85%;"><a href="/wiki/Fellow_of_the_Royal_Society_of_Edinburgh" class="mw-redirect" title="Fellow of the Royal Society of Edinburgh">FRSE</a></span></span> (8 November 1905 – 26 September 1975) was a British <a href="/wiki/Developmental_biologist" class="mw-redirect" title="Developmental biologist">developmental biologist</a>, <a href="/wiki/Paleontology" title="Paleontology">paleontologist</a>, <a href="/wiki/Geneticist" title="Geneticist">geneticist</a>, <a href="/wiki/Embryologist" class="mw-redirect" title="Embryologist">embryologist</a> and <a href="/wiki/Philosopher" class="mw-redirect" title="Philosopher">philosopher</a> who laid the foundations for <a href="/wiki/Systems_biology" title="Systems biology">systems biology</a>, <a href="/wiki/Epigenetics" title="Epigenetics">epigenetics</a>, and <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a>. </p><p>Although his theory of <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetic assimilation</a> had a <a href="/wiki/Darwinian" class="mw-redirect" title="Darwinian">Darwinian</a> explanation, leading <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biologists</a> including <a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a> and <a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> considered that Waddington was using genetic assimilation to support so-called <a href="/wiki/Lamarckian" class="mw-redirect" title="Lamarckian">Lamarckian</a> inheritance, the acquisition of inherited characteristics through the effects of the environment during an organism's lifetime. </p><p>Waddington had wide interests that included <a href="/wiki/Poetry" title="Poetry">poetry</a> and <a href="/wiki/Painting" title="Painting">painting</a>, as well as left-wing political leanings. In his book <i>The Scientific Attitude</i> (1941), he touched on political topics such as <a href="/wiki/Economic_planning" title="Economic planning">central planning</a>, and praised <a href="/wiki/Marxism" title="Marxism">Marxism</a> as a "profound scientific philosophy".<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Life">Life</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=1" title="Edit section: Life"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Conrad Waddington, known as "Wad" to his friends and "Con" to family, was born in <a href="/wiki/Evesham" title="Evesham">Evesham</a> to Hal and Mary Ellen (Warner) Waddington, on 8 November 1905. </p><p>His family moved to India and until nearly three years of age, Waddington lived in India, where his father worked on a tea estate in the <a href="/wiki/Wayanad_district" title="Wayanad district">Wayanad district</a> of <a href="/wiki/Kerala" title="Kerala">Kerala</a>. In 1910, at the age of four, he was sent to live with family in England including his aunt, uncle, and Quaker grandmother. His parents remained in India until 1928. During his childhood, he was particularly attached to a local druggist and distant relation, Dr. Doeg. Doeg, whom Waddington called "Grandpa", introduced Waddington to a wide range of sciences from chemistry to geology.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> During the year following the completion of his entrance exams to university, Waddington received an intense course in chemistry from <a href="/wiki/E._J._Holmyard" class="mw-redirect" title="E. J. Holmyard">E. J. Holmyard</a>. Aside from being "something of a genius of a [chemistry] teacher," Holmyard introduced Waddington to the "Alexandrian <a href="/wiki/Gnosticism" title="Gnosticism">Gnostics</a>" and the "Arabic <a href="/wiki/Alchemy" title="Alchemy">Alchemists</a>." From these lessons in metaphysics, Waddington first gained an appreciation for interconnected <a href="/wiki/Holistic" class="mw-redirect" title="Holistic">holistic</a> systems. Waddington reflected that this early education prepared him for <a href="/wiki/Alfred_North_Whitehead" title="Alfred North Whitehead">Alfred North Whitehead</a>'s philosophy in the 1920s and 30s and the <a href="/wiki/Cybernetics" title="Cybernetics">cybernetics</a> of <a href="/wiki/Norbert_Wiener" title="Norbert Wiener">Norbert Wiener</a> and others in the 1940s.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>He attended <a href="/wiki/Clifton_College" title="Clifton College">Clifton College</a> and <a href="/wiki/Sidney_Sussex_College,_Cambridge" title="Sidney Sussex College, Cambridge">Sidney Sussex College, Cambridge</a>. He took the Natural Sciences <a href="/wiki/Tripos" title="Tripos">Tripos</a>, earning a First in Part II in geology in 1926.<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> In 1928, he was awarded an Arnold Gerstenberg Studentship in the University of Cambridge, whose purpose was to promote "the study of Moral Philosophy and Metaphysics among students of Natural Science, both men and women."<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> He took up a Lecturership in Zoology and was a Fellow of <a href="/wiki/Christ%27s_College,_Cambridge" title="Christ's College, Cambridge">Christ's College</a> until 1942. His friends included <a href="/wiki/Gregory_Bateson" title="Gregory Bateson">Gregory Bateson</a>, <a href="/wiki/Walter_Gropius" title="Walter Gropius">Walter Gropius</a>, <a href="/wiki/C._P._Snow" title="C. P. Snow">C. P. Snow</a>, <a href="/wiki/Solly_Zuckerman" class="mw-redirect" title="Solly Zuckerman">Solly Zuckerman</a>, <a href="/wiki/Joseph_Needham" title="Joseph Needham">Joseph Needham</a>, and <a href="/wiki/John_Desmond_Bernal" class="mw-redirect" title="John Desmond Bernal">John Desmond Bernal</a>.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><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> His interests began with <a href="/wiki/Palaeontology" class="mw-redirect" title="Palaeontology">palaeontology</a> but moved on to the <a href="/wiki/Heredity" title="Heredity">heredity</a> and development of living things. He also studied philosophy. </p><p>During <a href="/wiki/World_War_II" title="World War II">World War II</a> he was involved in <a href="/wiki/Operational_research" class="mw-redirect" title="Operational research">operational research</a> with the <a href="/wiki/Royal_Air_Force" title="Royal Air Force">Royal Air Force</a> and became scientific advisor to the Commander in Chief of <a href="/wiki/RAF_Coastal_Command" title="RAF Coastal Command">Coastal Command</a> from 1944 to 1945. </p><p>After the war, in 1947, he replaced <a href="/wiki/Francis_Albert_Eley_Crew" title="Francis Albert Eley Crew">Francis Albert Eley Crew</a> as Professor of Animal Genetics at the <a href="/wiki/University_of_Edinburgh" title="University of Edinburgh">University of Edinburgh</a>.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> He would stay at Edinburgh for the rest of life with the exception of one year (1960–1961) when he was a Fellow on the faculty in the Center for Advanced Studies at <a href="/wiki/Wesleyan_University" title="Wesleyan University">Wesleyan University</a> in Middletown, Connecticut.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> His personal papers are largely kept at the University of Edinburgh library. </p><p>He died in Edinburgh on 26 September 1975. </p> <div class="mw-heading mw-heading2"><h2 id="Family">Family</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=2" title="Edit section: Family"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Waddington was married twice. His first marriage produced a son, C. Jake Waddington, professor of physics at the <a href="/wiki/University_of_Minnesota" title="University of Minnesota">University of Minnesota</a>, but ended in 1936. He then married architect <a href="/wiki/Margaret_Justin_Blanco_White" class="mw-redirect" title="Margaret Justin Blanco White">Margaret Justin Blanco White</a>, daughter of the writer <a href="/wiki/Amber_Reeves" title="Amber Reeves">Amber Reeves</a>, with whom he had two daughters, the anthropologist <a href="/wiki/Caroline_Humphrey" title="Caroline Humphrey">Caroline Humphrey</a> (1943–) and mathematician <a href="/wiki/Dusa_McDuff" title="Dusa McDuff">Dusa McDuff</a> (1945–).<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><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=3" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the early 1930s, Waddington and many other embryologists looked for the molecules that would induce the amphibian neural tube. The search was beyond the technology of that time, and most embryologists moved away from such deep problems. Waddington, however, came to the view that the answers to embryology lay in genetics, and in 1935 went to <a href="/wiki/Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Hunt Morgan's</a> <i>Drosophila</i> laboratory in California, even though this was a time when most embryologists felt that genes were unimportant and just played a role in minor phenomena such as eye colour. </p><p>In the late 1930s, Waddington produced formal models about how gene regulatory products could generate developmental phenomena, showed how the mechanisms underpinning <i>Drosophila</i> development could be studied through a systematic analysis of mutations that affected the development of the <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> wing.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> In a period of great creativity at the end of the 1930s, he also discovered mutations that affected cell phenotypes and wrote his first textbook of "developmental epigenetics", a term that then meant the external manifestation of genetic activity. </p><p>Waddington introduced the concept of <a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">canalisation</a>, the ability of an organism to produce the same <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> despite variation in genotype or environment. He also identified a mechanism called <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetic assimilation</a> which would allow an animal's response to an environmental stress to become a fixed part of its developmental repertoire, and then went on to show that the mechanism would work. </p><p>In 1972, Waddington founded the Centre for Human Ecology in the <a href="/wiki/University_of_Edinburgh" title="University of Edinburgh">University of Edinburgh</a>.<sup id="cite_ref-CHE_15-0" class="reference"><a href="#cite_note-CHE-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Epigenetic_landscape">Epigenetic landscape</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=4" title="Edit section: Epigenetic landscape"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Waddington's <a href="/wiki/Epigenetic_landscape" class="mw-redirect" title="Epigenetic landscape">epigenetic landscape</a> is a metaphor for how <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">gene regulation</a> modulates development.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Among other metaphors, Waddington asks us to imagine a number of marbles rolling down a hill.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The marbles will sample the grooves on the slope, and come to rest at the lowest points. These points represent the eventual <a href="/wiki/Biological_cell" class="mw-redirect" title="Biological cell">cell</a> fates, that is, <a href="/wiki/Biological_tissue" class="mw-redirect" title="Biological tissue">tissue</a> types. Waddington coined the term <a href="/wiki/Chreode" class="mw-redirect" title="Chreode">chreode</a> to represent this cellular developmental process. The idea was based on experiment: Waddington found that one effect of mutation (which could modulate the epigenetic landscape) was to affect how cells differentiated. He also showed how mutation could affect the landscape, and used this metaphor in his discussions on evolution—he emphasised (like <a href="/wiki/Ernst_Haeckel" title="Ernst Haeckel">Ernst Haeckel</a> before him) that evolution mainly occurred through mutations that affected developmental anatomy. </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_assimilation">Genetic assimilation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=5" title="Edit section: Genetic assimilation"><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:Lamarck_Compared_to_Darwin,_Baldwin,_Waddington.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg/550px-Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg.png" decoding="async" width="550" height="325" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg/825px-Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg/1100px-Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg.png 2x" data-file-width="1382" data-file-height="817" /></a><figcaption>Waddington's <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetic assimilation</a> compared to <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a>, <a href="/wiki/Darwinian_evolution" class="mw-redirect" title="Darwinian evolution">Darwinian evolution</a>, and the <a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a>. All the theories offer explanations of how organisms respond to a changed environment with adaptive inherited change.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></div> <p>Waddington proposed an evolutionary process, "<a href="/wiki/Genetic_assimilation" title="Genetic assimilation">genetic assimilation</a>", as a <a href="/wiki/Darwinian" class="mw-redirect" title="Darwinian">Darwinian</a> mechanism that allows certain acquired characteristic to become heritable. According to Navis, (2007) "Waddington focused his genetic assimilation work on the crossveinless trait of <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i>. This trait occurs with high frequency in heat-treated flies. After a few generations, the trait can be found in the population, without the application of heat, based on hidden genetic variation that Waddington asserted had been "assimilated".<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neo-Darwinism_versus_Lamarckism">Neo-Darwinism versus Lamarckism</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=6" title="Edit section: Neo-Darwinism versus Lamarckism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Waddington's theory of genetic assimilation was controversial. The <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biologists</a> <a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a> and <a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> both thought that Waddington was using genetic assimilation to support <a href="/wiki/Lamarckian" class="mw-redirect" title="Lamarckian">Lamarckian</a> inheritance. They denied that genetic assimilation had taken place, and asserted that Waddington had simply observed the <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> of genetic variants that already existed in the study population.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Other biologists such as <a href="/wiki/Wallace_Arthur" title="Wallace Arthur">Wallace Arthur</a> disagree, writing that "genetic assimilation, looks, but is not Lamarckian. It is a special case of the evolution of <a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">phenotypic plasticity</a>".<sup id="cite_ref-wallace-evo_21-0" class="reference"><a href="#cite_note-wallace-evo-21"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Adam S. Wilkins wrote that "[Waddington] in his lifetime... was widely perceived primarily as a critic of Neo-Darwinian evolutionary theory. His criticisms ... were focused on what he saw as unrealistic, 'atomistic' models of both gene selection and trait evolution." In particular, according to Wilkins, Waddington felt that the Neo-Darwinians badly neglected the phenomenon of extensive gene interactions and that the "randomness" of mutational effects, posited in the theory, was false.<sup id="cite_ref-Wilkins_2008_22-0" class="reference"><a href="#cite_note-Wilkins_2008-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> Even though Waddington became critical of the neo-darwinian synthetic theory of evolution, he still described himself as a Darwinian, and called for an <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">extended evolutionary synthesis</a> based on his research.<sup id="cite_ref-Wilkins_2008_22-1" class="reference"><a href="#cite_note-Wilkins_2008-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Reviewing the debate in 2015, the <a href="/wiki/Systems_biology" title="Systems biology">systems biologist</a> <a href="/wiki/Denis_Noble" title="Denis Noble">Denis Noble</a> writes however that </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>[Waddington] did not describe himself as a Lamarckian, but by revealing mechanisms of inheritance of acquired characteristics, I think he should be regarded as such. The reason he did not do so is that Lamarck could not have conceived of the processes that Waddington revealed. Incidentally, it is also true to say that Lamarck did not invent the idea of the inheritance of acquired characteristics. But, whether historically correct or not, we are stuck today with the term 'Lamarckian' for inheritance of a characteristic acquired through an environmental influence.<sup id="cite_ref-Noble2015_24-0" class="reference"><a href="#cite_note-Noble2015-24"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup></p></blockquote> <div class="mw-heading mw-heading2"><h2 id="As_an_organiser">As an organiser</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=7" title="Edit section: As an organiser"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Waddington was very active in advancing biology as a discipline. He contributed to a book on the role of the sciences in times of war, and helped set up several professional bodies representing biology as a discipline.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>A remarkable number of his contemporary colleagues in Edinburgh became Fellows of the Royal Society during his time there, or shortly thereafter.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Waddington was an old-fashioned intellectual who lived in both the arts and science milieus of the 1950s and wrote widely. His 1969 book <i>Behind Appearance; a Study of the Relations Between Painting and the Natural Sciences in This Century</i> (MIT press) not only has wonderful pictures but is still worth reading.<sup id="cite_ref-Ritterbush1970_27-0" class="reference"><a href="#cite_note-Ritterbush1970-27"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Waddington was, without doubt, the most original and important thinker about <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a> of the <a href="/wiki/Molecular_biology" title="Molecular biology">pre-molecular age</a> and the medal of the British Society for Developmental Biology is named after him.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p> Waddington co-founded The Institute for Advanced Studies in the Humanities at the University of Edinburgh in 1969 with Professor John MacQueen, Professor of Scottish Literature and Oral Tradition.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup></p><ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues,_1955.jpg" class="mw-file-description" title="Pages from a photograph album, given to Waddington by his colleagues on his 50th birthday."><img alt="Pages from a photograph album, given to Waddington by his colleagues on his 50th birthday." src="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg/120px-Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg" decoding="async" width="120" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg/180px-Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/84/Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg/240px-Page_from_a_photograph_album_given_to_C.H._Waddington_by_his_colleagues%2C_1955.jpg 2x" data-file-width="1536" data-file-height="1219" /></a></span></div> <div class="gallerytext">Pages from a photograph album, given to Waddington by his colleagues on his 50th birthday.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/120px-Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg" decoding="async" width="120" height="96" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/180px-Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/45/Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/240px-Page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg 2x" data-file-width="1536" data-file-height="1230" /></a></span></div> <div class="gallerytext"></div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/120px-Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg" decoding="async" width="120" height="97" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/180px-Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg/240px-Further_page_from_a_photograph_album_belonging_to_C.H._Waddington.jpg 2x" data-file-width="1536" data-file-height="1238" /></a></span></div> <div class="gallerytext"></div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="Selected_works">Selected works</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=8" title="Edit section: Selected works"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Books">Books</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=9" title="Edit section: Books"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Waddington, C. H. (1939). <i>An Introduction to Modern Genetics</i>. London : George Alien & Unwin.</li> <li>––– (1940). <i>Organisers & Genes</i>. Cambridge: Cambridge University Press.</li> <li>––– and others <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 book cs1"><a rel="nofollow" class="external text" href="https://archive.org/details/in.ernet.dli.2015.206560/page/n7/mode/1up"><i>Science and Ethics</i></a>. George Allen & Unwin. 1942 – via <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Science+and+Ethics&rft.pub=George+Allen+%26+Unwin&rft.date=1942&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fin.ernet.dli.2015.206560%2Fpage%2Fn7%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></li> <li>––– (1946). <i>How Animals Develop</i>. London : George Allen & Unwin.</li> <li>––– <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a rel="nofollow" class="external text" href="https://archive.org/details/rkas.2138.scientificattitu0000chwa/page/n1/mode/1up"><i>The Scientific Attitude</i></a> (2nd ed.). Pelican Books. 1948 – via <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Scientific+Attitude&rft.edition=2nd&rft.pub=Pelican+Books&rft.date=1948&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Frkas.2138.scientificattitu0000chwa%2Fpage%2Fn1%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></li> <li>––– (1953). <i>The Epigenetics of birds</i>. Cambridge : Cambridge University Press.</li> <li>––– (1956). <i>Principles of Embryology</i>. London : George Allen & Unwin.</li> <li>––– (1957). <i>The Strategy of the Genes</i>. London : George Allen & Unwin.</li> <li>––– (1959). <i>Biological Organisation Cellular and Subcellular : Proceedings of a Symposium</i>. London: Pergamon Press.</li> <li>––– (1960). <i>The Ethical Animal</i>. London : George Allen & Unwin.</li> <li>––– (1961). <i>The Human Evolutionary System</i>. In: Michael Banton (Ed.), <i>Darwinism and the Study of Society</i>. London: Tavistock.</li> <li>––– (1961). <i>The Nature of Life</i>. London : George, Allen, & Unwin.</li> <li>––– (1962). <i>New Patterns in Genetics and Development</i>. New York: Columbia University Press.</li> <li>––– (1966). <i>Principles of Development and Differentiation</i>. New York: Macmillan Company.</li> <li>––– (1970). 72). <i>Behind Appearance : A Study in the Relationship Between Painting and the Natural Sciences in this Century</i>. The MIT Press.</li> <li>–––, ed. (1968–72). <i>Towards a Theoretical Biology</i>. 4 vols. Edinburgh: Edinburgh University Press.</li> <li>–––, <a href="/wiki/Anthony_Kenny" title="Anthony Kenny">Kenny, A.</a>, <a href="/wiki/H._Christopher_Longuet-Higgins" class="mw-redirect" title="H. Christopher Longuet-Higgins">Longuet-Higgins, H.C.</a>, <a href="/wiki/John_Lucas_(philosopher)" title="John Lucas (philosopher)">Lucas, J.R.</a> (1972). <i>The Nature of Mind</i>, Edinburgh: Edinburgh University Press (1971-3 <a href="/wiki/Gifford_Lectures" title="Gifford Lectures">Gifford Lectures</a> in Edinburgh, <a rel="nofollow" class="external text" href="https://www.giffordlectures.org/lectures/nature-mind">online</a>)</li> <li>–––, Kenny, A., Longuet-Higgins, H. C., Lucas, J. R. (1973). <i>The Development of Mind</i>, Edinburgh: Edinburgh University Press (1971-3 Gifford Lectures in Edinburgh, <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080621235816/http://www.giffordlectures.org/Browse.asp?PubID=TPTDOM&Cover=TRUE">online</a>)</li> <li>––– (1973) <i>O.R. in World War 2: Operational Research Against the U-Boat</i>. London: Elek Science.</li> <li>–––, & Jantsch, E. (Eds.). (1976). (published posthumously). <i>Evolution and Consciousness: Human Systems in Transition</i>. Addison-Wesley.</li> <li>––– (1977) (published posthumously). <i>Tools for Thought</i>. London: Jonathan Cape.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Papers">Papers</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=10" title="Edit section: Papers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Waddington, C. H. (1942). Canalization of development and the inheritance of acquired characters. <i>Nature</i> 150 (3811):563–565.</li> <li>--- (1946). Human Ideals and Human Progress. <i>World Review</i> August:29-36.</li> <li>––– & Carter T. C. (1952). Malformations in mouse embryos induced by trypan blue. <i>Nature</i> 169 (4288):27-28.</li> <li>––– (1952). Selection of the Genetic Basis for an Acquired Character. <i>Nature</i> 169 (4294):278.</li> <li>––– (1953). Genetic assimilation of an acquired character. <i>Evolution</i> 7:118–126.</li> <li>––– (1953). Epigenetics and evolution. <i>Symposia of the Society of Experimental Biology</i> 7:186–199.</li> <li>––– (1956). Genetic assimilation of the bithorax phenotype. <i>Evolution</i> 10:1–13.</li> <li>––– (1961). Genetic assimilation. <i>Advances in Genetics</i> 10:257–290.</li> <li>––– (1974). A Catastrophe Theory of Evolution. <i>Annals of the New York Academy of Sciences</i> 231:32–42.</li> <li>––– (1977).(published posthumously). Whitehead and Modern Science. <i>Mind in Nature: The Interface of Science and Philosophy</i>. Ed. John B. Cobb and David R. Griffin. University Press of America.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=11" title="Edit section: Notes"><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-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text">This was the essence of the approach that won the 1995 Nobel prize in medicine for <a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a> and <a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a><sup id="cite_ref-Winchester_12-0" class="reference"><a href="#cite_note-Winchester-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=C._H._Waddington&action=edit&section=12" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWaddington,_C._H.1948" class="citation book cs1">Waddington, C. H. (1948). <a rel="nofollow" class="external text" href="https://archive.org/details/rkas.2138.scientificattitu0000chwa/page/n1/mode/1up"><i>The Scientific Attitude</i></a> (2nd ed.). Pelican Books. p. <a rel="nofollow" class="external text" href="https://archive.org/details/rkas.2138.scientificattitu0000chwa/page/104/mode/1up">104</a> – via <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Scientific+Attitude&rft.pages=104&rft.edition=2nd&rft.pub=Pelican+Books&rft.date=1948&rft.au=Waddington%2C+C.+H.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Frkas.2138.scientificattitu0000chwa%2Fpage%2Fn1%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text">Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905–26 September 1975." <i>Biographical Memoirs of Fellows of the Royal Society</i> <b>23</b>, 575-622. pp. 575-76. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsbm.1977.0022">10.1098/rsbm.1977.0022</a></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Waddington, C. H. 1975. <i>The Evolution of an Evolutionist</i>. Ithaca, NY: Cornell University Press. Pg. 2.</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 – 26 September 1975." <i>Biographical Memoirs of Fellows of the Royal Society</i> <b>23</b>, 575-622. Pg 577.</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><i>Supplement, Historical Register of the University of Cambridge, 1921-30</i>, Cambridge: Cambridge University Press, 1932, p. 63.</span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text">Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 – 26 September 1975." <i>Biographical Memoirs of Fellows of the Royal Society</i> <b>23</b>, 575-622. Pp. 579-580.</span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text">Yoxen, Edward. 1986. "Form and Strategy in Biology: Reflections on the Career of C. H. Waddington." In <i>A History of Embryology</i>, edited by T. J Horder, J. A Witkowski, and C. C Wylie. Cambridge: Cambridge University Press. pp. 310-11.</span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</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="http://blogs.royalsociety.org/history-of-science/2013/07/08/good-fellowship/">"Good Fellowship - The Repository - Royal Society"</a>. <i>blogs.royalsociety.org</i><span class="reference-accessdate">. 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Retrieved <span class="nowrap">4 April</span> 2010</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Guide+to+the+Center+for+Advanced+Studies+Records%2C+1958+-+1969&rft.pub=Wesleyan.edu&rft_id=http%3A%2F%2Fwww.wesleyan.edu%2Flibr%2Fschome%2FFAs%2Fce1000-137.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-10">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDarling2019" class="citation web cs1">Darling, Elizabeth (2019). <a rel="nofollow" class="external text" href="https://www.oxforddnb.com/display/10.1093/odnb/9780198614128.001.0001/odnb-9780198614128-e-112261">"White [married name Waddington], (Margaret) Justin Blanco (1911–2001), architect"</a>. <i>Oxford Dictionary of National Biography</i>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fodnb%2F9780198614128.013.112261">10.1093/odnb/9780198614128.013.112261</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-861412-8" title="Special:BookSources/978-0-19-861412-8"><bdi>978-0-19-861412-8</bdi></a><span class="reference-accessdate">. Retrieved <span class="nowrap">18 January</span> 2024</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Oxford+Dictionary+of+National+Biography&rft.atitle=White+%5Bmarried+name+Waddington%5D%2C+%28Margaret%29+Justin+Blanco+%281911%E2%80%932001%29%2C+architect&rft.date=2019&rft_id=info%3Adoi%2F10.1093%2Fodnb%2F9780198614128.013.112261&rft.isbn=978-0-19-861412-8&rft.aulast=Darling&rft.aufirst=Elizabeth&rft_id=https%3A%2F%2Fwww.oxforddnb.com%2Fdisplay%2F10.1093%2Fodnb%2F9780198614128.001.0001%2Fodnb-9780198614128-e-112261&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text">Robertson, Alan. 1977. Conrad Hal Waddington. 8 November 1905 – 26 September 1975. <i>Biographical Memoirs of Fellows of the Royal Society</i> <b>23</b>, 575-622. P. 578</span> </li> <li id="cite_note-Winchester-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-Winchester_12-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWinchester2004" class="citation journal cs1">Winchester, Guil (2004). <a rel="nofollow" class="external text" href="http://www.cell.com/current-biology/pdf/S0960-9822(04)00683-9.pdf">"Edward B. Lewis 1918-2004"</a> <span class="cs1-format">(PDF)</span>. <i>Current Biology</i>. <b>14</b> (18) (published 21 September 2004): R740–742. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2004CBio...14.R740W">2004CBio...14.R740W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2004.09.007">10.1016/j.cub.2004.09.007</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/15380080">15380080</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:32648995">32648995</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Biology&rft.atitle=Edward+B.+Lewis+1918-2004&rft.volume=14&rft.issue=18&rft.pages=R740-742&rft.date=2004&rft_id=info%3Adoi%2F10.1016%2Fj.cub.2004.09.007&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A32648995%23id-name%3DS2CID&rft_id=info%3Apmid%2F15380080&rft_id=info%3Abibcode%2F2004CBio...14.R740W&rft.aulast=Winchester&rft.aufirst=Guil&rft_id=http%3A%2F%2Fwww.cell.com%2Fcurrent-biology%2Fpdf%2FS0960-9822%2804%2900683-9.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</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://web.archive.org/web/20161013223611/https://www.ibiology.org/ibiomagazine/eric-wieschaus-and-christiane-nusselin-volhard.html">"Eric Wieschaus and Christiane Nüsslein-Volhard: Collaborating to Find Developmental Genes"</a>. iBiology. 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Retrieved <span class="nowrap">9 November</span> 2016</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Conrad+Waddington&rft.pub=Centre+for+Human+Ecology&rft_id=http%3A%2F%2Fwww.che.ac.uk%2Fwhat-we-do%2Fconrad-waddington%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text">Goldberg, A. D., Allis, C. D., & <a href="/wiki/Emily_Bernstein" title="Emily Bernstein">Bernstein, E.</a> (2007). Epigenetics: A landscape takes shape. Cell, 128, 635-638.</span> </li> <li id="cite_note-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-17">^</a></b></span> <span class="reference-text"><a rel="nofollow" class="external text" href="https://www.academia.edu/14699065/Compelled_by_the_Diagram_Thinking_through_C._H._Waddington_s_Epigenetic_Landscape_Contemporaneity_August_2015_,">Allen, Matthew. 2015. "Compelled by the Diagram: Thinking through C. H. Waddington’s Epigenetic Landscape." <i>Contemporaneity</i> <b>4</b>.</a></span> </li> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNavis2007" class="citation encyclopaedia cs1">Navis, Adam R. (2007). 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BioEssays&rft.atitle=The+molecular+and+mathematical+basis+of+Waddington%27s+epigenetic+landscape%3A+A+framework+for+post-Darwinian+biology%3F&rft.volume=34&rft.issue=2&rft.pages=149-157&rft.date=2012&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19632484%23id-name%3DS2CID&rft_id=info%3Apmid%2F22102361&rft_id=info%3Adoi%2F10.1002%2Fbies.201100031&rft.aulast=Huang&rft.aufirst=Sui&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" class="Z3988"></span></span> </li> <li id="cite_note-Noble2015-24"><span class="mw-cite-backlink"><b><a href="#cite_ref-Noble2015_24-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNoble2015" class="citation journal cs1"><a href="/wiki/Denis_Noble" title="Denis Noble">Noble, Denis</a> (2015). <a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.120071">"Conrad Waddington and the origin of epigenetics"</a>. <i>Journal of Experimental Biology</i>. <b>218</b> (6): 816–818. <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/10.1242%2Fjeb.120071">10.1242/jeb.120071</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/25788723">25788723</a>.</cite><span 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Retrieved <span class="nowrap">24 July</span> 2020</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=IASH+at+50&rft.pub=University+of+Edinburgh&rft_id=https%3A%2F%2Fwww.iash.ed.ac.uk%2F50&rfr_id=info%3Asid%2Fen.wikipedia.org%3AC.+H.+Waddington" 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=C._H._Waddington&action=edit&section=13" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20030504104717/http://www.nahste.ac.uk/pers/w/GB_0237_NAHSTE_P0355/">NAHSTE Project Record of C.H. Waddington</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20051125091507/http://zygote.swarthmore.edu/gene3a.html">Induction and the Origin of Developmental Genetics - works by Salome Gluecksohn-Schoenhimer and Conrad Hal Waddington</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20060514181106/http://www.epigeneticsnews.com/">Epigenetics News</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 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.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 href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a 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 class="mw-selflink selflink">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"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q721128#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q721128#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a 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href="https://isni.org/isni/0000000108705484">ISNI</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://viaf.org/viaf/12383955">VIAF</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://id.worldcat.org/fast/6298/">FAST</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.oclc.org/worldcat/entity/E39PBJyRCJMmBQvdwmDGjDqxXd">WorldCat</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-even" 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/120829088">Germany</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/n50019537">United States</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb12353465d">France</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb12353465d">BnF data</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00459952">Japan</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=skuk0005772&CON_LNG=ENG">Czech Republic</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://data.bibliotheken.nl/id/thes/p068367481">Netherlands</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://authority.bibsys.no/authority/rest/authorities/html/90111857">Norway</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://lod.nl.go.kr/resource/KAC199629024">Korea</a></span></li><li><span class="uid"><a 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class="external text" href="https://www.leopoldina.org/mitgliederverzeichnis/mitglieder/member/Member/show/conrad-hal-waddington/">Leopoldina</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">People</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://trove.nla.gov.au/people/1020794">Trove</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://www.deutsche-biographie.de/pnd120829088.html?language=en">Deutsche Biographie</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://www.deutsche-digitale-bibliothek.de/person/gnd/120829088">DDB</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" 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